Front Cover
 Title Page
 Front Matter
 Table of Contents

The scarab beetles of Florida (Coleoptera: Scarabaeidae)
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00000088/00001
 Material Information
Title: The scarab beetles of Florida (Coleoptera: Scarabaeidae)
Series Title: Arthropods of Florida and Neighboring Land Areas
Physical Description: v. : illus. ; 27 cm.
Language: English
Creator: Woodruff, Robert E
Florida -- Division of Plant Industry
Publisher: Florida Dept. of Agriculture and Consumer Services, Division of Plant Industry
Place of Publication: Gainesville
Publication Date: 1973
Subjects / Keywords: Scarabaeidae   ( lcsh )
Insects -- Florida   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: v. 1, p. 182-200.
Additional Physical Form: Electronic version available on the World Wide Web as part of the Linking Florida's Natural Heritage Collection.
Statement of Responsibility: by Robert Eugene Woodruff.
 Record Information
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0021
notis - AAA6418
alephbibnum - 000005307
oclc - 01092408
lccn - 74622007
Classification: lcc - QL434 .A75 v.8
ddc - 595/.2/09759 s|595.7/64
System ID: UF00000088:00001


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Table of Contents
    Front Cover
        Cover 1
        Cover 2
    Title Page
        Page i
    Front Matter
        Page ii
    Table of Contents
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Full Text

And Neighboring Land Areas




Taxonomic Entomologist

Fig. 1 Phanaeus igneus floridanus d'Olsoufieff (male)
(actual size = 1 8mm)

AND CONSUMER SERVICES Halwin L. Jones, Director
Post Office Box 1269
Doyle Conner, Commissioner Gainesville, Florida 32601

And Neighboring Land Areas




Taxonomic Entomologist

Doyle Conner, Commissioner
Halwin L. Jones, Director

Post Office Box 1269
Gainesville, Florida 32601

Contribution No. 260 Bureau of Entomology '

Release Date: September 1, 1973



Vernon Conner, Chairman (Citrus) .......................................... ..... ........ M ount Dora

Roy Vandegrift, Jr., Vice Chairman (Vegetables) ....................................... .. . Canal Point

Colin English, Sr. (Citizen at-Large) ................................................... . Tallahassee

Lawrence W . Clements (Citrus) ........................................ ................... . Bartow

Joseph Welker (Ornamental Horticulture) ............................................... Jacksonville

Fred J. W esemeyer (Commercial Flower) ................................................... Ft. Myers

Foster Shi Smith (Forestry) ............................................. ............. . Starke

Felix H. Uzzell (Apiary) ......................................................... . . Sebring

Halwin L. Jones, Secretary ............................................. ............... . Gainesville


Halwin L. Jones, Director .............................................. ............... . Gainesville

S. A. Alfieri, Jr., Assistant Director .. .................................................. Gainesville

G. D. Bridges, Chief, Bureau of Citrus Budwood Registration ................................ Winter Haven

J. K. Condo, Chief, Bureau of Plant Inspection ......................... ... .......... . .. Gainesville

H. A. Denmark, Chief, Bureau of Entomology ................................ ........ . Gainesville

P. M. Packard, Chief, Bureau of Apiary Inspection ....................................... . Gainesville

C. Poucher, Chief, Bureau of Pest Eradication and Control ................................. Winter Haven

C. P. Seymour, Chief, Bureau of Plant Pathology ........................................ Gainesville

A. L. Taylor, Chief, Bureau of Nematology ............... ......................... ...... Gainesville

This public document was promulgated at a cost of $6994.95 or $2.33 per copy. It
makes available to all interested persons the results of arthropod faunal studies emphasizing
Florida and the Circum-Caribbean Region.



Title Page ...... - --- ------------ ----
Division of Plant Industry Technical Council & Administrative Staff .-- .----------
Table of Contents ---..... ------------.-...-----------------

List of Figures ----..... ------------- - ------------------ -
Foreword ..- -------------------------------
Preface -------------------------------------------- ---------------------------

Acknowledgm ents -. ------------------ ---------------------------------------------------- ------------------------
Abstract --- ----------------------------
Introduction ----...- . ---- ... .. ------. ------ --...-- -------------------------------------------- --

H historical R esum e - . -------....... ....------------ --- ----------- -- --- -- -------
Collecting Techniques -- --- - ----. ---- -- ---- - ----------------
Form at of Presentation ----------- --. --------------------------------------------
Florida Biogeography -- ....------- --- - - ----------------- ----

' eral Account of the Family
Taxonomy .--------.. --....
S Morphology ........- -...--....
Biology -----
S Behavior .........----------..-------
Diel Flight Activity ......
Associations ...------
Nidification -----
S Sound Production ......
Orientation .-----------
S Pasture Ecosystems ...-----
Parasites .. .. --
Zoogeography .-................
S Paleontology ..-..'..... ..-- -
Economic Importance .....-----

Preliminary Checklist of Florida Scarabaeidae ---...-- ------------------

- Systematic Account --.-.---.--- .....................-----------------------.
Key to the Florida Subfamilies of Scarabaeidae .-..............-- .....---..-..---------------------..
Subfamily Scarabaeinae .....----. ..---------------------------------------------------
Key to the Florida Tribes and Genera of Scarabaeinae ..-..........-- ........ ............--- ...
Tribe Scarabaeini .- ....------------.. . ...... --------.-. ---- -------------...------...
Genus Deltochilum .-.. ---------------------------------------...
Genus Canthon .................................. ...--
Genus Boreocanthon .......--...--....------------...- ..... ......---...-----.-----------------
Genus Melanocanthon --- -------.....----------....
Genus Glaphyrocanthon ............--------------- -----.. ---------------------
Genus Pseudocanthon .......-.........................-.-................... -. ......
Tribe Coprini .............-- ........----- ---.--.------..-- ...........--------------------..
Genus Ateuchus ------- ----- ...........-----------------........-
Genus Dichotomius ..- - ..............-------------..--------........ ------------------------
G enus Copris .---.-- . ---. -...........- ..........----- .. -------..---...-... ....-----.-
Genus Phanaeus ..............------ --------------------- --------------------------------









--------------------------------- --------------------- ----------------------------------------------------------
------------------------------------------------------------------------------ -----------------------------------
------------------------------------------------------------- ---------- -----------------------------------------
---------------- --------------------------------------------------------- ----------------------------------------
------------------------------------- ------------------------------------ ----------------------------------------
-------------------------- ---------------------------------- ----------------------------------------------
------------ - ------------------------------------------------------------ ----------------------------------------
--------------------- --------------------------------------------------------------------- -------------------
----------------------- --------------------------------------------------- -----------------------------------------
---------------------- I ---------------------------------- ----------- ----------------------------------------
---------------------------------------------- I --------------------------------------------------------------------


Tribe Onthophagini - ...... ..................... ...................
Genus Onthophagus ..... - ...- ........---------------..-..-........-. .. ..
T ribe O niticellini ---------------....------------------------------.......---------......... ...........................
Genus Oniticellus ---. - --- ----.... ................. . . ..... . . .... .-- ........
Subfam ily A phodiinae ---------------------------- --- -------......................
Key to the Florida Genera of Aphodiinae -----..-...--- -.......
Genus Aphodius --- .......... . -......... .............................................
Genus M yrmecaphodius -.. - -..... -----.. --- ..----.- ......-.............. . . . .
Genus Euparia ..............---------------..--------....... ...-....----.-- ----- ---....... ---............ ............
Genus Ataenius - --........~~-- -..... .........--- - -------.. ............ ......
Genus Pseudataenius ---------.. . ..............--------------------------.......................
Genus Aphotaenius --..--.-----...----------......----------..--.. ------.........--...- ....
Genus Psam m odius .....---------.. .............. ... .-.. ..... ......... ...........................
Genus Pleurophorus .......------- --------------............

Genus Rhyssemus --------------------- ..........
Subfamily Hybosorinae -. - - -- - ..... ..... -..................
Genus H ybosorus .. . ..... . ..... ........ ..............................
Subfamily Ochodaeinae ---. --. - .... --- .................
Genus O chodaeus .... .......... ............................................
Subfam ily Geotrupinae .-----.......-. ----------..- .. ..-.....--.......-- ......---........
Key to the Florida Tribes and Genera of Geotrupinae --------- ------------
Tribe Bolboceratini .-- .- -- --- ............................. ..
Genus Bolboceras .........................- ........ ---- ........ ................................
Genus Bradycinetulus ---- -----------. ------------ --....................
Genus Bolbocerosoma ------ -... .. . .. ..... ....................
Genus Eucanthus ....-........ ~_~.........-----------. -- -.. ------......... ....................
Tribe Geotrupini --- ---- - ...-----.... .... ...... .........................
Genus Geotrupes ----- ---------. ........... ... ............ ..
G enus Peltotrupes - ....... ......... ...........................................
Genus Mlycotrupes . ........ ... .......... ....... ..........................
Subfam ily Acanthocerinae .-..... ---------------. ---------..... .-..- ...............
Key to the Florida Genera and species of Acanthocerinae ---.-- ....---................
Genus Acanthocerus ... ------ ------------------...................................
Genus Cloeotus - ------.... .. . .... ..............................
Bibliography ---- ---- -----..-- .. .........................................
A ppendices -.-----. ----...... .--------------------------.... . -----..-..-... ...... -.... ---.. ......---.......- ........-... -.. -.......-... ......... ................
Appendicesx ....-----------------.. -.................. ,.............




Phanaeus igneus floridanus, habitus drawing .............................. .............. Cover
Abdominal spiracles of Pleurosticti and Laparosticti . ............................. ....... 1
Bait traps in wooded area .. . ....................................................... 2
"Push up" of Peltotrupes ................................................ .............. 3
Habitat at Cape Sable, Florida ........................................................ 3

8. Beach area at Cape Sable, Florida ..... .............................................


Geotrupes egeriei and pupal cell ...........
Dichotomius carolinus larva and cell .......
Currents of the Gulfstream ...............
Map of Florida, average summer rains .....
Map of Florida, average winter rains .......
Map of Florida, average July temperature ...
Map of Florida, average January temperature
Map of Florida, hours of frost per year .....
Map of Florida, general soil distribution ....
Map of Florida, average annual rainfall .....

20. Habitat along Tamiami Trail near Ochopee ............................
21. Habitat along Apalachicola River at Torreya State Park ...............
22. Habitat in dunes at St. Andrews State Park .............. ........
23. Rosemary bush (Ceratiola ericoides) in dunes .........................
24. Habitat of Mycotrupes cartwrighti in Tallahassee "Red Hills" ..........
25-29. Strategus antaeus, larval spiracles ..................... ...............
30. Diagram of scarab relationships based on follicle structure ...............
31-32. Polyphylla occidentalis, elytral scales ............................ .
33-38. Polyphylla occidentalis, antenna ................. .....................
39-58. Key subfamily and generic taxonomic characters .......................
59. Typical scarab larva ........................................
60-62. Morphological characters used in larval taxonomy .......................
63. Typical scarab pupa ... .......... ..............................
64. Strategus antaeus, larval epipharynx .............. . ..................
65-66. Strategus antaeus, stridulatory area of larval mandible ..................
67-68. Distribution map of gopher tortoise ................................
69-70. Packrat droppings at Florida Caverns St. Pk. .......................
71-72. Packrat nest on Key Largo ........................................
73. Distribution map of Florida packrats ................................
74. Pocket gopher mounds ........................................
75. Distribution map of Florida pocket gophers ............................
76-77. Imported fire ant nests ... . ........ ..............................


.... . . . . . . . . . . .. . . . 8
............ .. . .. . 10
.................... 10
.................... 11
............... . . . 11
............... . . . . 11
............. . . . . . 11-12
............... . . . 12
............... . . . 13
.............. . . 14-15
.............. . . 15-16
.................... 17
............... . . . 17
.................... 18
............... . . . . 18
............... . . . 19
......... ...... . . . 19


Audiospectrograph of scarab sounds .. ................................................ . 22
Florida burrowing owl at its burrow ............................ .... ................. 23
Burrowing owl pellets with scarab remains ................................................ 23
Deltochilum g. gibbosum, habitus drawing ............................................ . 34
Deltochilum g. gibbosum, distribution map ................................................. 34
Canthon pilularius, habitus drawing ........................... ........................ 35
Canthon spp., taxonomic characters ................ .......................... 37
Canthon spp., distribution maps .................................................. . 38,40
Boreocanthon depressipennis, habitus drawing .................. ..... ................. 41
Boreocanthon spp., distribution maps .... ......... ............... ..................... 41-42
Characters distinguishing Canthon from Boreocanthon ....................................... 42
Melanocanthon punctaticollis, habitus drawing ............................................ 43


. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .�

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .�

� .

. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . ..
. . . . . . .. .. .. .. ..
..... . .... .. .. ..
. . . . . . . . . . . . . . . . .


Fig. Pages
108-110. Melanocanthon granulifer, stereoscan photos ........................................ ... .... 43
111-112. Melanocanthon spp., distribution maps ..................................................... 44
113-114. M elanocanthon spp., genitalia drawings ................................................. . 45
115-118. Melanocanthon spp., distribution maps .............................................. . . 45-46
119-121. Melanocanthon punctaticollis, stereoscan photos ............................................. 46
122. Glaphyrocanthon v. viridis, habitus drawing .............................................. 47
123-124. Glaphyrocanthon v. viridis, distribution maps .............................................. 47
125. Pseudocanthon perplexus, habitus drawing ................................................. 48
126-127. Pseudocanthon perplexus, distribution maps ................................................ 49
128. Ateuchus lecontei, habitus drawing ....................................................... 50
129-130. Ateuchus spp., taxonomic characters ........................................ ............. 51
131-132. Ateuchus lecontei, distribution maps ...................................................... 52
133. Dichotomius carolinus, habitus drawing ................................................... 53
134-135. Dichotomius carolinus, distribution maps .................................................. 54
136. Copris minutus, habitus drawing ........................................... ............. 55
137-138. Copris spp., taxonomic characters ......................................................... 56
139-144. Copris spp., distribution maps ................................................... . 56, 57, 59
145. Phanaeus vindex, lateral view of male .................................................... 60
146. Phanaeus vindex, stereoscan photo of elytron ............................................... 61
147-148. Phanaeus igneus igneus, stereoscan photo of elytron ......................................... 61
149-150. Phanaeus igneus floridanus, stereoscan photo of elytron ....................................... 62
151-154. Phanaeus spp., distribution maps ....................................... ................ 63
155. Onthophagus p. polyphemi, habitus drawing ............................................. . 64
156. Onthophagus hecate blatchleyi, pronotal horn ............................................... . 67
157-176. Onthophagus spp., distribution maps ................................................... 67-76
177. Oniticellus cubiensis, habitus drawing ..................................................... 77
178. Aphodius fimetarius, habitus drawing ........................................ ........... 80
179-180. Aphodius spp., taxonomic characters ................................................ ..... 83
181-211. Aphodius spp., distribution maps ..................................................... 84-100
212. Myrmecaphodius excavaticollis, habitus drawing : ............ . ........................... 101
213. Myrmecaphodius excavaticollis, adult in ant nest ........................................ 102
214. Myrmecaphodius excavaticollis larva in ant nest ........................................... 102
215-217. Myrmecaphodius excavaticollis, stereoscan photos ...................................... 102-103
218-219. Myrmecaphodius excavaticollis, distribution maps .......................................... 103
220. Euparia castanea, habitus drawing ....................................................... 104
221. Euparia castanea, distribution maps ........................ ......... ............... 105
222. Ataenius saramari, habitus drawing .................................................... . 106
223-231. Ataenius spp., taxonomic characters .................................................. 107-108
232-234. Ataenius spp., distribution maps ................................................... . 111-112
235-237. Ataenius alternatus, stereoscan photos .................................................... 112
238-245. Ataenius spp., distribution maps ................................................... . 114-116
246-248. Ataenius havanensis, stereoscan photos ................................................... 11'
249-260. Ataenius spp., distribution maps ................................................... . 117-125
261-263. Ataenius miamii, stereoscan photos ...................................................... 125
264-267. Ataenius spp., distribution maps .................................................. . 123-124
268-272. Ataenius picinus, stereoscan photos ................. ................................ ......125-126
273-274. Ataenius platensis, distribution maps ..................................... ....... . 126
275-277. Ataenius platensis, stereoscan photos ..................................................... 127
278-280. Ataenius spp., distribution maps ................................................... . 128-129








Hybosorus illigeri, taxonomic characters ............................................... 147
Ochodaeus frontalis, habitus drawing ............... ................................... 148
Ochodaeus spp., taxonomic characters .. . ......................................... 149-151
Ochodaeus frontalis, distribution maps .. ............ ............................. . 151
Bolboceras floridensis, habitus drawing .... ............................................ 154
Bolboceras floridensis, distribution maps .............. . ................................. 155
Bradycinetulus ferrugineus, habitus drawing .............. . .. ... ..... ............. . . . .. 155
Bradycinetulus ferrugineus, distribution maps ................... ....................... 156
Bolbocerosoma hamatum, habitus drawing .......................................... 157
Bolbocerosoma hamatum, distribution maps ........... ............... ............... . 157
Eucanthus subtropicus, habitus drawing .......................... . ................ 158
Eucanthus spp., stereoscan photos ............... ...................................... 159
Eucanthus spp., distribution maps .. ................................................ 160-161
Geotrupes egeriei, habitus drawing .. ................................................ . 162
Geotrupes spp., taxonomic characters .. . ......................................... 163
Geotrupes spp., distribution maps .............. ....................... . . .... . 164-165
Peltotrupes profundus, habitus drawing ................................... .... ........ 165
Peltotrupes profundus, photo of pair in copula ..................... .................. 166
Peltotrupes profundus, male and female anterior tibiae ..................................... 166
Peltotrupes spp., distribution maps ............... .. ................. ................. . 166
Peltotrupes young, habitat photos .. ................................................ . 167
Mycotrupes gaigei, habitus drawing .............. ................................ 169
Mycotrupes spp., distribution maps ................ . ................................... 171
Acanthocerus aeneus, habitus drawing ..... ............................... ... . . . 174
Acanthocerinae larval characters .. . ............................................... . 175
Acanthocerus aeneus, distribution maps ............... ................................ 176
Cloeotus globosus, habitus drawing ................ . ................. ................. 177
Acanthocerinae, adult taxonomic characters ...................................... ..... 177-178
Cloeotus globosus, stereoscan photo .. . .......... ................................. . 179
Cloeotus aphodioides, distribution maps .................................................. 179
Cloeotus globosus, stereoscan photo ............................................. . 179-181
Cloeotus globosus, distribution maps ...................................................... 181

Ataenius rudellus, stereoscan photos ................
Ataenius spp., distribution maps ...................
Pseudataenius socialis, habitus drawing ............
Pseudataenius socialis, distribution map ............
Aphotaenius carolinus, habitus drawing .............
Psammodius malkini, habitus drawing ..............
Psammodius malkini, stereoscan photos .............
Psammodius spp., distribution maps ................
Pleurophorus longulus, habitus drawing .............
Pleurophorus longulus, distribution maps ............
Rhyssemus scaber, habitus drawing ................
Hybosorus illigeri, habitus drawing ................
Hybosorus illigeri, distribution maps ...............

...................................... 129
.................................. 130-132
................ ...................... 134
................ ...................... 135
................ ...................... 136
...................................... 137
........................... . . . . . 138-139
.................................. 139-141
..................................... . 142
...................................... 143
................ ...................... 144
...................................... 146
...................................... 146


Beetles of the family Scarabaeidae are both destruc-
tive and beneficial. Many, like the Japanese beetle
and Asiatic garden beetle, along with the larvae or
"white grubs", cause millions of dollars damage an-
nually. Others are economically important because they
are intermediate hosts for parasites of domestic ani-
mals. Most of the dung beetles are important elements
in the pasture ecosystem where they annually break
down tons of animal dung. By doing so they incor-
porate much of it into the soil, increasing fertility, and
at the same time destroying the habitat for the larvae
of many pest flies.
This group of beetles is one of the largest in the
animal kingdom, represented by perhaps 30,000 species.
Scarabs have been subjects of interest throughout re-
corded history. They were worshipped by the ancient
Egyptians, and their images are found in precious
stones of both ancient and modern jewelry. Many of
the early naturalists (e.g., Fabre) found their behavior
unique and fascinating, but it is still poorly under-
stood today. Few generalizations can be made, because
of the specific nature of the biology, ecology, and
behavior of each species.
The present faunal study is provided as a manual
to assist in the identification of the Florida species.
The specific identity of an organism is paramount to
an understanding of the role it plays in the environ-
ment. The name is the "key" to the published litera-
ture, and an absolute must before controls are attempt-
ed. It is unfortunate that we do not have adequate
guides for insect identification as we do with birds,
mammals, fish, reptiles, and amphibians. There are
inherent difficulties because of insects' small size; but
the primary reason for the lack of such identification
manuals is that thorough faunal studies have not been
conducted on much of our planet. Dr. Woodruff's
manual is based on an extensive survey over a 15 year
period and on the examination of over 1 million speci-
mens. He brings together all the existing information
about each species, provides drawings and photographs,
and presents keys for the identification of the Florida
Dr. Woodruff was born on 20 July 1933 at Kennard,
Ohio. He has been interested in the natural sciences
since childhood, and he entered the Junior Science
Fairs of the Ohio Academy of Science, in 1950 and
1951, receiving superior awards both years and a schol-
arship to any of the 10 state universities. He enrolled
at Wabash College (Crawfordsville, Ind.) in 1951, and
the following year he transferred to Ohio State Uni-
versity from which he received the B.S. degree in 1956.
From 1952 to 1955 he was an assistant in the Dept.
of Natural History of the Ohio State Museum where
he gained valuable experience and training under Dr.
Edward S. Thomas and Mr. Robert Goslin. From 1955
to 1957 he held a graduate assistantship in the Dept.
of Zoology and Entomology at Ohio State University,
under Prof. J. N. Knull. It was this experience which
led him to specialization on the beetle family Scara-
baeidae. Prof. Knull introduced him to the "scratch

board" technique of beetle drawings which he has
used in most of his publications and for the 32 habitus
drawings presented herein.
From 1957 to 1958 he was employed as a medical
entomologist with the Kentucky State Health Dept.
(Louisville) working on St. Louis Encephalitis and
related mosquito projects. He attended special courses
of instruction on insects of medical importance from
the U. S. Public Health Service, Communicable Disease
Center, Atlanta, and from the Tennessee Valley Au-
thority, Wilson Dam, Alabama.
In March 1958 he joined the staff of entomologists
at the State Plant Board of Florida (now Division of
Plant Industry). He was initially employed as the
"Survey Entomologist" to coordinate the federal-state
Cooperative Economic Insect Survey which he ful-
filled until 1963. His taxonomic responsibilities are
for the insect orders Orthoptera and Coleoptera. His
current duties include the identification and curatorial
responsibilities for these orders and the development
of the entomological portion of the DPI library.
In September 1963 he entered the Graduate School
of the University of Florida, from which he received
the PhD degree in 1967. As a part of his studies he
attended a session of the Organization for Tropical
Studies at the University of Costa Rica. His research
for the dissertation culminated in the original version
of the present study. In the subsequent 5 years much
study and collecting has resulted in more than doubling
the data on Florida scarabs.
His research has taken him to much of the U. S.
and the following countries where he has collected
and studied specimens: Argentina, Australia, Bolivia,
Brazil, Colombia, Costa Rica, Cuba, El Salvador,
Guatemala, Honduras, Jamaica, Mexico, Nicaragua,
Peru, and Venezuela.
He has attended more than 20 national and inter-
national scientific meetings, including the recent 14th
International Congress of Entomology in Canberra,
Australia. He has presented papers at meetings of the
Entomological Society of America, Florida Entomologi-
cal Society, National Pest Control Association, Ohio
Academy of Sciences, and Sociedad Mexicana de
He is a member of the Association for Tropical
Biology, Coleopterists Society, Entomological Society
of America, Florida Entomological Society, Gamma
Sigma Delta, Phi Kappa Psi, Sigma Xi, Sociedad
Mexicana de Entomologia, and Society of Systematic
He has been a Research Associate in Natural
Sciences of the Florida State Museum since 1962. He
was recently elected to the Board of Directors of the
"North American Beetle Fauna Project." He has served
as merit badge counsellor for the Boy Scouts in all
Natural History subjects. He has received grants from
the Ohio Academy of Sciences, United States Public
Health Service, National Science Foundation, Smith-
sonian Institution, United States Department of Agri-
culture, Australian Academy of Sciences, and the

Florida State Museum. In 1971 he attended the Sum-
mer Institute for Systematics at the Smithsonian In-
stitution. He is listed in American Men of Science,
Directory of Coleoptera Collections, Directory of Zoo-
logical Taxonomists, International Scholars Directory,
Personalities of the South, and Who's Who in the South
and Southwest.
He served as Editor for the "Journal of the Newell
Entomological Society" when he was President in 1965-
66. He was Editor of the "Coleopterists Newsletter"
in 1970. He has been Associate Editor of the "Florida
Entomologist" since 1969. He has been Editor of the
"Coleopterists Bulletin" since 1971. He was recently
appointed as Managing Editor of "Insect World
He has published over 80 scientific papers, most of
which deal with his primary research interests on the

systematics, biology, and ecology of the beetle family
Scarabaeidae, with special emphasis on dung beetles
and those inquilines associated with ants and ter-
mites. Recently his design was chosen in competition
as the symbol for the XV International Congress of
Entomology to be held in Washington, D. C. in 1976.

Harold A. Denmark
Chief of Entomology
Division of Plant Industry
Florida Department of Agriculture
and Consumer Services


This study began in 1958 when I joined the staff
of the Division of Plant Industry, Florida Department
of Agriculture and Consumer Services (formerly Flor-
ida State Plant Board). My interest in the family
Scarabaeidae dates back to 1954 when I started a
similar study of the fauna of Ohio. Most of the earlier
years were devoted to extensive collecting in order to
become acquainted with the species and to obtain
distributional data. Only in the past few years has
this information been organized into the present study.
The original manuscript for this study was submitted
in 1967 as a dissertation to the Graduate Council of
the University of Florida in partial fulfillment of the
requirements for the degree of Doctor of Philosophy.
The general format remains the same, although the
basic information has been up-dated and the number
of records has more than tripled.
Faunal studies are fairly complete for most of the
vertebrate animals, but they are woefully lacking for
most invertebrate groups. In Florida there have been
several noteworthy exceptions in the insects: Odonata
of Florida (Byers, 1930), Mayflies of Florida (Berner,
1950), Water Beetles of Florida (Young, 1954), and
the previous volumes in the present series [e.g., Lepi-
doptera of Florida (Kimball, 1965); Armored Scales
of Florida (Dekle, 1965)]. Other similar volumes are
in preparation as a part of this series on the Arthropods
of Florida.
Herbert Osborne, one of our greatest pioneer en-
tomologists, once said (1912:63): "While the prepara-
tion of such [faunal] lists may by some be considered
as a rather easy part of entomological investigations,
it appears to me that accurately done work of this
kind [faunal studies] becomes of the highest scientific
value, and that we may very well encourage it to the
greatest extent possible." In a paper presented to the
Florida Academy of Sciences on the opportunities for
research in Florida, Kurz (1937:8) stated that ". . . it
becomes readily apparent that we need local or regional
florass' and faunass' by which naturalists can readily
and with certainty identify species of particular inter-
est." His statement is equally applicable 35 years later.
Our ecological problems and the fragility of our
environment, of which the general public has recently
become aware, has long been well-known to natural
history students. I am convinced that faunal and floral
studies, verified by permanently preserved museum
specimens, are just as important a part of "environ-
mentalism" as anti-pollution campaigns. We should at
least know what we have destroyed!
Florida is an ideal state for faunal studies because
it is a peninsula and is delimited on three sides. It
is an important area from a zoogeographic standpoint
because of the proximity of the West Indies and be-
cause of the high percentage of endemic species. A
symposium on Floridian-Antillean zoogeography, held
at the 1960 meetings of the Entomological Society of
America, suggested that only a few groups of insects
had been sufficiently studied in both areas to draw
general conclusions. This study should help fill that

The present study is not merely taxonomic, nor is
it ecological, biological, ethological, or zoogeographical;
it is faunal in nature, encompassing all of these disci-
plines. It is a compilation of all existing information,
including both published and original data. It is prob-
ably based on a greater number of specimens of this
family than have been studied from any other state in
the U. S., or in fact any comparable geographic area
in the New World. It should be primarily useful to
those who wish to identify Florida specimens and then
to determine what is known about a particular species.
If it serves this purpose and stimulates additional
studies, I will be adequately rewarded for the efforts
involved in its preparation.


As an entomologist with the Division of Plant
Industry, Florida Department of Agriculture and Con-
sumer Services, I have been able to pursue several
aspects of this work over a period of 15 years. For
their encouragement and understanding I thank the
following administrators of this organization: The
Honorable Doyle E. Conner, Commissioner of Agricul-
ture; H. L. Jones and the late W. G. Cowperthwaite,
Directors, Division of Plant Industry; and H. A. Den-
mark, Chief, Bureau of Entomology.
I have also had the benefit of close association
with my colleagues on the staff of the Division of
Plant Industry: G. W. Dekle, F. W. Mead, and H. V.
Weems, Jr. They have provided specimens, advice,
and companionship on many field trips.
In addition to support from the Florida Department
of Agriculture, I have received several grants which
provided funds for travel, technicians, and general as-
sistance from: 1) the National Science Foundation
(for a summer traineeship through the Organization
for Tropical Studies in Costa Rica and for participa-
tion in the Summer Institute for Systematics at the
Smithsonian Institution); 2) the University of Florida
(for field work in Jamaica through National Institute
of Health grants and for study of the scarab inquiline
associated with the imported fire ant in the United
States and South America, through USDA grants);
and 3) the Australian Academy of Sciences, the Com-
monwealth Scientific and Industrial Research Organiza-
tion, the University of Florida, and Dr. H. E. Hinton
for aid in attending the International Congress of
Entomology in Australia, and for field work in
I am indebted to several museums for the loan of
specimens and for the use of their facilities during
personal visits. Letters in parentheses in the following
list are abbreviations used in the text when citing
material; the name of the curator(s) who provided
assistance is listed after the abbrevation: Academy of
Natural Sciences of Philadelphia (ANSP), S. S. Ro-
back; American Museum of Natural History (AMNH),
M. A. Cazier, Patricia Vaurie; Canadian National Col-

election (CNC), H. F. Howden; Chicago Natural History
Museum (CNHM), R. L. Wenzel, H. S. Dybas; *Florida
State Collections of Arthropods (FSCA), H. V. Weems,
Jr.; Museum of Comparative Zoology (MCZ), P. J.
Darlington, J. F. Lawrence; North Carolina State Uni-
versity (NCS), T. B. Mitchell; Ohio State University
(OSU), J. N. Knull, F. J. Moore, C. A. Triplehorn;
Purdue University, Blatchley Collection (PU), Leland
Chandler; United States National Museum (USNM),
O. L. Cartwright, R. D. Gordon, P. J. Spangler; Uni-
versity of Florida (UF), T. J. Walker, L. C. Kuitert;
University of Miami (UM), H. F. Strohecker; Uni-
versity of Michigan Museum of Zoology (UMMZ),
R. D. Alexander, and T. H. Hubbell.
I am also indebted to the following individuals
who loaned or donated material from their private
collections: Ross H. Arnett, Jr., Bernard Benesh, L. J.
Bottimer, O. L. Cartwright, Neil Chernoff, B. K. Dozier,
C. A. Frost, D. H. Habeck, E. I. Hazard, H. F. Howden,
J. W. McReynolds, F. J. Moore, D. R. Paulson, Joe
Schuh, W. R. Suter, P. A. Thomas, and D. W. Thornton.
The following scarab specialists provided assistance
in many ways, including checking specimens, providing
loans, and supplying data, literature, and specimens:
B. Benesh, W. J. Brown, O. L. Cartwright, R. W.
Dawson, T. Fincher, R. D. Gordon, G. Halffter, A. R.
Hardy, H. E. Hinton, H. F. Howden, A. Martinez,
E. Matthews, P. Reyes C., P. O. Ritcher, M. W. San-
derson, and Patricia Vaurie.
Light traps were one of the greatest sources of
material for this study. Samples were obtained from a
network of stations throughout the state with the aid
of the following individuals: W. W. Baker, A. H. Boike,
P. E. Briggs, R. E. Brown, S. H. Brown, F. A. Bu-
chanan, T. W. Boyd, F. S. Blanton, E. M. Collins, Jr.,
H. W. Collins, L. Collins, E. E. Crooks, G. W. Desin,
C. F. Dowling, H. M. Faircloth, E. H. Frederic, J. C.
Hanlon, D. L. Harris, E. I. Hazard, L. A. Hetrick,
J. Hayward, E. W. Holder, Jr., E. G. Kelsheimer, R. L.
King, J. H. Knowles, M. Lutrick, D. L. Mays, E. S.

*The Florida State Collection

Mercer, R. T. McMillan, E. P. Merkel, M. L. Messec,
T. Morris, A. L. O'Berry, J. W. Patton, J. W. Perry,
A. M. Phillips, W. H. Pierce, J. E. Porter, W. C.
Rhoades, R. W. Swanson, W. B. Tappan, L. W. Taylor,
J. W. Wilson, D. P. Wojcik, and D. O. Wolfenbarger.
The greatest number of specimens, from a single
location, was received from Tall Timbers Research
Station, Leon County, Florida. I take pleasure in thank-
ing the Tall Timbers Foundation and particularly the
following personnel for various favors: E. V. and Roy
Komarek, W. W. Baker, D. L. Harris, Leroy Collins,
W. H. Whitcomb, Awinash Bhatkar, and R. H.
Arnett, Jr.
I wish to thank a great many individuals, espe-
cially inspectors of the Division of Plant Industry and
U. S. Department of Agriculture, for providing assist-
ance and specimens. Space does not permit a full list-
ing here, but their names are included in the appendix
as collectors. For photographic assistance I am indebted
to the Division of Plant Industry photographers, E. M.
Collins, Jr., Mildred Eaddy, Jerry Messec, and E. L.
Wells. I wish to thank the following librarians of
the Division of Plant Industry for assistance in locat-
ing references: Marguerite Batey, Louise Henley, Irene
Ayres, and Andrew Kolesar. I am grateful to W. A.
Auffenberg, Curator of Natural Sciences, Florida State
Museum, for providing unpublished data for the dis-
tribution map of the gopher tortoise in Florida. P. S.
Callahan and Patricia Carlisle (USDA, Basic Behavior
Lab, Gainesville) provided assistance in the form of
stereoscan photos. For assistance in nearly all aspects
of the work, I thank my technician, Brenda Beck, and
my secretary, Irene Ayres.
I wish to thank the members of my Ph.D. super-
visory committee for their assistance: W. G. Eden,
E. S. Ford, D. H. Habeck, L. A. Hetrick, and T. J.
Walker, Jr.
Last, but not least, I wish to thank my wife, Evelyn,
for her constant encouragement and great care and
perseverance in typing the manuscript.

of Arthropods is composed

of several collections which were previously maintained as
separate: Univ. Fla., Agr. Exp. Sta.; Univ. Fla., Dept.
Entomology; Florida State Museum; Florida State Plant
Board; and Division of Plant Industry. My private collec-
tion (REW) of Scarabaeidae is located with this collection
which is housed by the Division of Plant Industry, Florida
Dept. Agr., Gainesville, Fla. 32601.

In this faunal study 248 species and subspecies of Scarabaeidae are recorded from Florida. Only the Laparosticti,
representing 115 species and subspecies, are treated in detail, although a checklist is provided for the entire family.
Keys are presented for the identification of all taxonomic units, with many of the morphological characters illustrated
by line drawings or stereoscan photographs. Habitus drawings are presented for one representative of each of the 32
genera of Florida Laparosticti.
Data are presented for each species in the following format: synonymy, diagnosis, taxonomic notes, distribution
and zoogeography, biology, specimens examined, and selected references. Distribution maps, showing detailed Florida
records and the general range, are provided for all species. Data from well over 1 million specimens are presented
in tabular form in the appendices. The bibliography contains 643 references. Numerous habitat photographs are
The following taxonomic changes are made: Aphodius geomysi Cartwright is synonymized under Aphodius
aegrotus Horn; Peltotrupes profundus dubius Howden is synonymized under Peltotrupes profundus Howden; Phanaeus
index cyanellus Robinson and Phanaeus difformis magnificens Robinson are synonymized under Phanaeus index


The family Scarabaeidae is one of the largest
families of beetles in the world. Arnett (1966:166)
estimated the number of species at 18,000, although
this is probably a conservative figure. In America north
of Mexico there are approximately 1,400 described
species, of which only about 600 were recognized
in 1910 (Blatchley, 1910:910). Thus the number has
more than doubled in the past 60 years.
The taxonomy of the U. S. species is relatively
stable-that is, few species are described each year.
Several taxonomists (e.g., Cartwright, Dawson, Gordon,
Hardy, Howden, Matthews, Sanderson, Vaurie, and
myself) are actively engaged in revising certain por-
tions of the family. Numerous generic revisions, during
the past 15 years, have clarified many of the problems,
but much remains to be done. Many genera of Melo-
lonthinae and Rutelinae have been so incompletely
studied that it will be several years before their tax-
onomy is on a par with the better studied groups.
Although taxonomically the family is one of the
better known of the larger families, the literature is
voluminous, scattered, and often difficult to secure.
The family has never been monographed for North
America or the U.S., and as Arnett (1962:410) stated,
"A comprehensive survey of this group is badly need-
ed." Such an undertaking would be a worthwhile but
S extremely time consuming proposition, and such an
opus is not imminent. I therefore believe that local
or regional faunal studies such as this will help fill
this void.
Many of the gaps in our knowledge become espe-
cially apparent after a detailed local faunal study. And
unfortunately, because of the large number of species
involved, it has not been possible to pursue and clarify
many of the problems encountered. However, I have
tried to point them out in the species discussions,
hoping they will stimulate future work along these
The family is one of the most diverse in the order
Coleoptera, not only in morphology, but in biology,
ecology, and behavior. On the basis of morphology
and habits it has been divided into two groups-the
Laparosticti and the Pleurosticti. The former was char-
acterized by having the abdominal spiracles situated
in a line on the membrane between the sternites and
tergites (Fig. 3), and included the dung-feeding and
scavenger species. These are represented in Florida by
the following six sub-families: Scarabaeinae, Aphodii-
nae, Geotrupinae, Hybosorinae, Ochodaeinae, and
Acanthocerinae. The Pleurosticti were characterized by
having most of the abdominal spiracles situated on the
upper portion of the sternites (Fig. 2, 4), and included
primarily the plant feeders and chafers. These are rep-
resented in Florida by the following four subfamilies:
Melolonthinae, Rutelinae, Cetoniinae, and Dynastinae.
Although current concepts indicate that the division
into Laparosticti and Pleurosticti is probably untenable
(Ritcher: 1969a), they are used here for convenience
(see discussion under morphology).
Due to limitations of time and space I have treated
the family in two parts, the first of which deals only
with the Laparosticti. However, data have been accu-

Fig. 2-4. Spiracle location in the Laparosticti and Pleurosticti:
2) Dorsal view of the abdomen of Euphoria sepulchralis
(Fab.) with the elytra and wings removed. Note the charac-
teristic positions of the spiracles found in the Pleurosticti.
3) Dorsal view of the abdomen of Copris minutus (Drury)
with elytra and wings removed. Note the characteristic posi-
tion of the spiracles found in the Laparosticti. 4) Lateral view
of the abdomen of Euphoria sepulchralis (Fab.).

mulated for the Pleurosticti during this study, and a
similar volume is in preparation on that group. A
preliminary checklist of the Florida species in both
groups is included.
Any faunal study obviously will not be up-to-date
when it is published. Specimens are constantly being
added to the collections which will modify the state-
ments made under the species discussions. Nearly all
material on hand has been recorded, except in the
subfamily Aphodiinae. In this subfamily over 1 million
specimens have been sorted from light trap samples
and await processing. The time required to sort, iden-
tify, and record this number would unduly delay com-
pletion of the manuscript, and more material would
accumulate in the meantime.
There have been scattered records of Florida Scara-
baeidae since the beginning of binomial nomenclature
(Linnaeus, 1758). Aside from the descriptions of new
species, the first list of Florida species was that of
Schwarz (1878). In this list of Florida Coleoptera,
based on previous records and 12 locations visited by
the author, he recorded 84 species of Scarabaeidae.
Schaupp (1878) added four species to this list. Horn
(1880) recorded a single species in his treatise on the
Coleoptera of the Florida Keys. Hubbard (1894)
described three new dung feeding species in his classic
study of the guests of the Florida land tortoise. Slosson
(1893) recorded one species from Pensacola and eight
species from Suwannee Springs. Hamilton (1894) re-
corded 11 species from Lake Worth and added 6
species later (1895). Slosson (1895) added two species
to the Lake Worth list. Castle and Laurent (1896)
listed 14 species from the vicinity of Enterprise. Wick-
ham (1909) reported 26 species from 7 Florida

Blatchley, between 1902 and 1927, published
numerous notes on the Florida fauna, culminating in
his "Scarabaeidae of Florida" (1927-30). In this
study he listed 194 species and subspecies for the
state, although he included all records regardless of
their validity. More will be said about this paper later.
C. C. Goff studied the arthropods in the burrows of
the Florida pocket gopher and gopher tortoise. In the
first he found five scarabs, three of which were un-
described (Hubbell and Goff, 1939), and in the latter
he recorded seven scarabs (Young and Goff, 1939).
Young and Thames (1949) listed 42 species of
Phyllophaga, and Dawson (1952 & 1967) recorded 15
species of Serica. Frost (1964) recorded 23 species
from blacklight traps at Archbold Biological Station
(Highlands Co.). In the present study I have recorded
248 species and subspecies, excluding all previous
erroneous records but including 11 questionable spe-
cies (indicated by an asterisk on the checklist).
There have been only three extensive faunal studies
of the family Scarabaeidae in portions of the U. S.
These are: Blatchely (1910) in his "Coleoptera of
Indiana"; Dawson (1922), the "Scarabaeidae of Ne-
braska"; and Blatchley (1927-30), the "Scarabaeidae
of Florida." The latter was less comprehensive than
the other two, containing no keys to separate the
species nor any illustrations. It was based on most
of the available collections at the time, but the State
had been poorly surveyed. One of the major advances
in collecting, the blacklight trap, has come about since
that study.
The diversity of food, habitats, and behavior of the
many species requires a similar diversity in collecting
techniques. Nearly every habitat, except aquatic, has
some species of Scarabaeidae, and the various tech-
niques must be tried in each of these habitats to secure
the maximum number of specimens and species.
Most of the species of Laparosticti are coprophagous
or necrophagous, and, by searching for such foods, one
can often obtain large numbers of some species. Cow
dung is one of the most universally present foods,
and it hosts many species. However, a few species
appear to be found only on the dung of one animal
(e.g., pocket gopher, gopher tortoise, deer). It is some-
times difficult to locate dung of such wild animals,
and most of the beetles associated with these hosts
remain rare in collections. Although some species com-
plete their life-cycle in dung or carrion, many species
bury the food beneath the source (e.g., Copris spp.,
Phanaeus spp.) or transport it some distance away
(e.g., Canthon spp.). It is therefore often necessary
to dig beneath the food source or search the immediate
vicinity for specimens.
Trapping offers an excellent method for securing
specimens of certain species, with a minimum amount
of effort from the collector. The food material (e.g.,
dung, carrion, etc.) can be used with some success,
but often artificial baits are more readily obtained and
easier to use. The bait traps, which I have used, con-
sisted of any convenient container ,(usually tin cans

or pint glass jars) sunk into the ground, level with
the top (Fig. 5). The soil was packed tightly around
the rim to permit easy access by the beetles. In the
case of liquid baits, the trap was about half filled
with bait. The traps were normally checked about
every two days, the bait strained through a tea
strainer, returned to the trap, and the trap reburied.
The specimens were washed in water to remove the
bait and dirt and then were transferred to 70% iso-
propyl alcohol.
Liquid baits (containing a small amount of the
attractant in water) that produced specimens include
proprionic acid, butyric acid, asafoetida, amyl acetate,
and fermenting solutions of malt, molasses, and yeast.
For certain species, combinations of the above proved
more effective (e.g. malt and proprionic acid for some
Geotrupinae). Preliminary testing suggested that a
bait consisting of one teaspoon of powdered yeast in
one pint of water is effective for several species. In
the case of malt and molasses, the bait appeared to
be more effective after fermentation had begun; usually
within two days after mixing. Bait trapping (especially
with malt) was the only convenient method for collect-
ing large numbers of many burrowing groups for
which the food habits are unknown (e.g., Mycotrupes,
Peltotrupes). The use of specific attractants offers
a fertile field for future investigation.

- ~-i45;

Fig. 5. Bait traps in place in a wooded area. Such traps,
using fermenting malt as bait, are especially useful in collect-
ing Bolboceras, Geotrupes, Mycotrupes, and Peltotrupes.
Matthews (1972:6) found that the most effective
bait in Australia was marsupial entrails. He stated
that: "It may be surmised that the beetles respond
most strongly to complex organic molecules of elaborate
structure, and least strongly to simple decomposition
products such as scatole or proprionic acid. Excrement
while still in the intestine has undergone least decom-
position and therefore contains the highest proportion
of complex molecules. The adaptive significance of
this response is obvious, as it enables the beetles to
locate the food with the highest energy content. For
this reason it is impossible to substitute any artificial

baits for excrement or entrails, as the complex sub-
stances needed, even if they could easily be obtained,
would decompose too rapidly."
Many of the Scarabaeidae are attracted to light,
and various designs of light traps are effective in
collecting specimens. Blacklight (actually ultraviolet)
is much more effective than white or visible light. The
traps which I have utilized are modifications of those
originally designed by the USDA for European chafer
(Amphimallon majalis Raz.) surveys. Specimens were
collected in 70% isopropyl alcohol placed in the col-
lecting container. For a discussion of the nature of
light sources, trap designs, and additional information
on light trapping, the reader is referred to the follow-
ing papers: USDA (1961); Hollingsworth, Hartsock,
and Stanley (1963); Frost (1952, 1958, 1963, 1964,
1966). The advent of the blacklight has nearly revolu-
tionized collecting in this group of beetles and has
resulted in more specimens and records than all other
methods combined. The ecology and ethology are still
poorly known for many species attracted to blacklight,
but this attractant has provided much useful seasonal
and geographical distribution data in addition to abun-
dant specimens for morphological study.
Other mass collecting methods used were modified
Berlese or Tulgren funnels and Malaise-type traps.
Some of the larger species (e.g., Deltochilum, Dichoto-
mius) have been trapped occasionally in mist nets
used for birds and bats.
Specimens were often floated from cow dung, leaf
litter, plant roots, etc., by placing the material in a
pail of water. Digging was often the only known resort
for collecting some species, especially those which
were found in the burrows of pocket gophers and the
gopher tortoise. The Geotrupinae often burrow to
great depths (e.g., 6 to 10 ft. in Mycotrupes and
Peltotrupes. These burrows are usually marked by a
large "push up" of soil at the entrance (Fig. 6), and
they can be followed by inserting a straw into the
hole and digging along side. Fortunately for the

Fig. 6. Typical "push-up" marking the burrow of Peltotrupes
profundus Howden. The burrows often exceed six feet in
depth. (Photo by Alvah Peterson)

collector these deep burrows are usually vertical.
Several species, especially Aphodiinae, occur in leaf
mold or pine needles and were found by scarping away
the surface litter and inspecting the soil surface.
Many of these also were taken in the Berlese funnel.
The genus Psammodius often occurs near beaches, and

Fig. 7. General view of the habitat at Cape Sable, Fla.
(Everglades National Park). Note the dominance of cacti,
yucca, and agave.

Fig. 8. The beach area at Cape Sable, Fla. Scarabs found
in such habitats include Alaenius rudellus Fall, Ataenius
miamii Cartwr., and Psammodius spp.

specimens were found by pulling up plants in such
habitats (Fig. 8). They were sometimes found under
boards and the windows of debris behind the high tide
Nearly all specimens were collected into 70%
isopropyl alcohol which permitted easy genitalic dis-
sections later. Specimens can be preserved indefinitely
in this solution and are available for future mounting
or dissection. The immature stages often were found

Fig. 9. Geotrupes egeriei Germar after just emerging from
the pupal cell composed of cow dung.

in dung or in the soil and usually were reared easily
in salve tins. They were killed in boiling water and
preserved in 70% isopropyl alcohol.

Keys:-Dichotomous keys are presented for all of
the taxonomic units (subfamily, tribe, genus, species)
found in Florida. The keys to species within a genus
follow the general discussion of the genus. In cases
where there is a single species, it is keyed out with
the genus in the key to genera of each subfamily or
tribe. All of the keys are reversible, with numbers
in parentheses indicating the rubric to consult for
retracing one's steps. The keys are artificial in the
sense that they use morphological structures which are
easily seen or by which specimens are easily sep-
arated, without regard to any phylogenetic importance
or scheme. Most of the keys are original, but where
they are not, the source is indicated. Free use has
been made of existing keys, but they have been modi-
fied for Florida forms and will not necessarily apply
to other regions. Every effort has been made to make
them as clear, concise, and easy to use as possible.
They are all dichotomous, with the couplets reading
parallel. References are made to illustrations wherever
Generic treatment.-The genera are arranged as
nearly as possible in phylogenetic order. The following
standard format is used for each genus: 1) Reference
is made to a figure which illustrates one species of
the genus; in most cases a typical species, but not
necessarily the type species. 2) Synonymy: includes
only the different names, including misspellings and
nomina nuda, under which the genus has been placed.
The citations include only the original use of the name
and not subsequent usage. 3) Listing of the type
species (formerly called genotype, but not to be con-
fused with this term in genetics); by whom and how
designated (e.g., monotypy, subsequent designation,

Fig. 10. Third instar larva of Dichotomius carolinus (L.).
Note the "hump-backed" appearance characteristic of many
dung beetle larvae.

Fig. 11. Dung cell containing a mature larva of Dichotomius
carolinus (L.). Note the area around the opening which is
being repaired by the larva.

etc.). 4) Diagnosis: usually a short synopsis of the
most distinctive features of the genus, but confined to
characters applicable to Florida species. 5) Taxonomic
Notes: discussion of the synonymy; listing of sub-
genera; status of the classification (e.g., recently
revised, in need of revision, etc.); variation; subspecies.
6) Distribution and Zoogeography: general zoogeo-
graphic region occupied (e.g., Neotropical, Nearctic,


Fig. 12. Currents of the Gulf Stream. Such patterns are sig-
nificant in interpreting the zoogeography and possible origin
of the Florida fauna.

etc.); number of species in the world, U. S., and
Florida; discussion of introduced species. 7) Biology:
food habits of adults and larvae; ecological notes
(e.g., limiting factors such as soil type, climate, verte-
brate associations); behavior, including nidification,
mating, and stridulation; information on the life cycle;
indication of the immature stages known and a brief
characterization of the larvae of the genus. 8) Selected
References: includes author, date, and page citation
to the most important papers on the genus; complete
citations are presented in the bibliography. 9) Key
to the Florida species of the genus.
Species treatment.-The species are arranged under
each genus in alphabetical order. The information on
each species is presented in the following format: 1)
Synonymy: includes all different combinations of names
under which the species has been known. Only the
original citation for each combination is cited, but
additional information on synonymy is discussed under
"Taxonomic Notes." During the course of preparing
the synonymy, a complete list of all known references
was prepared, but it was not included in the manu-
script due to space limitations. 2) Diagnosis: a brief
resume of key features separating the species from
other Florida species. In many cases reference is made
to characters distinguishing it from closely related
c species, even though they are not fouhd in Florida.

It was felt that complete morphological descriptions
were not appropriate for this study and would unduly
lengthen the manuscript. The latest generic revision
can be consulted for such complete descriptions. Ref-
erence is made to drawings where pertinent. 3)
Taxonomic Notes: the status of the species and rea-
sons for any synonymy are explained. Variation,
subspeciation, and relationships are discussed. 4)
Distribution and Zoogeography: reference is made to
a distribution map for each species, except for a few
species with single records or for those that were
discovered too late and for which fragmentary infor-
mation was known (e.g., some of the Ataenius). In-
cluded is a detailed map of the Florida records and
a generalized map showing the broader distribution in
the New World. The previous literature records for
Florida are listed and discussed. Both literature rec-
ords and personal records are plotted on the map.
When only county records are known, they are plotted
in the center of the county (this can be determined
by reference to the data listed in the appendices).
Any questionable records are discussed, and the speci-
mens which I have seen are listed either in the section
under "Specimens Examined" or in the Appendices.
The generalized distribution also is based on literature
records and on specimens I have examined. In many
cases the records are not sufficient to present a true

picture of the distribution, but I feel that any map
is better than none. The data I have summarized here
should form a basis for future detailed studies. Inade-
quate as they may be, these are the only maps pub-
lished for a great number of the species. 5) Biology:
this is essentially a "cacth all" category for nearly
all other aspects of the species. It includes data on
abundance, seasonal distribution, food habits, habitat
preferences, collecting notes, nidification, stridulation,
and any inquilinous relationships. Notation is made
of whether the immature stages are known, and if so,
they are briefly characterized. 6) Specimens Examined:
includes the total number of specimens personally
examined, the number of Florida localities represented
and the number of collection records. If less than 10
records are known, the label data is included here;
if there are more than 10 records, these data are pre-
sented in the appendices (to which a reference is
made). Abbreviations for the various museums and col-
lections are listed under the acknowledgments. 7)
Selected References: these include all the pertinent
taxonomic and biological references, not repeating
those listed in the synonymy under each species. Ab-
breviated citations (author, date, page) only appear
here. Only those papers which contain significant data
are included, and catalogues, or state lists are generally
excluded. Complete citations are included in the ter-
minal bibliography.
Some new species (discovered during this study)
are listed here by number only, with a brief notation
about them, pending formal description elsewhere.
They are not included in the keys at present. They
are listed at the end of each genus, after the alpha-
betical arrangement of described species, so the reader
will be aware of their existence.

Studies on a diversity of plant and animal groups

36 - 32 - .- 1


Fig. 13. Average summer rains in inches (from Atlas of
Florida). *


Fig.14. Average winter rains in inches (from Atlas of Florida).

have indicated that peninsular Florida has been an
important area for the evolution and dispersion of the
flora and fauna of the southeastern Coastal Plain.
This situation is due to many factors, among which
are the geographical position, the physical shape, and
the geological history of the state. Not only was the
peninsula a refugium for certain species during glacial
periods, but its unique geographical position and geo-
logical history appear to have been conducive to the
establishment of groups of endemic, closely allied,
allopatric species of apparently recent origin (Hubbell,
There is little doubt about the importance of the
Florida peninsula as a refugium and an area with a
high percentage of endemic forms. Many of the relict
species occupy the sandhills and sand-pine scrub as
defined by Laessle (1958). It appears that these dry
areas often represent fossil dunes or prehistoric shore-
lines. They are now arranged in a pattern of faunal
and floral islands throughout the state, characterized
by well drained soils, often surrounded by swampy
areas of lower elevation. The age of the marine ter-
races is not firmly established, but several of those
previously considered of Pleistocene age are probably
Pliocene or Late Miocene (Alt and Brooks, 1965).
Small (1929), Blatchley (1932), and Barbour
(1945) have all discussed the general natural history
of southern Florida and the precariousness of the
habitats and their associated endemic faunas. Mc-
Cluney and others (1971) pointed out how rapidly
the environmental destruction is taking place. In many
cases the fauna already has been severely affected. In
a review of this book (Woodruff, 1971) I stated:
"There is probably no area of comparable size in the
U.S. which is more likely to have its flora and fauna
depleted than south Florida."
It is also apparent that our present knowledge
about the distribution of most insects, including the
Scarabaeidae, is too fragmentary to draw many sig-
nificant conclusions. In most cases the known distri-

Fig. 15. Average July temperature (from Atlas of Florida).


Fig. 16. Average January temperature (from Atlas of Florida).

butional patterns are based on such meager data that
they cannot be interpreted with any degree of confi-
dence. This situation became very apparent when I
tried to analyze the data on the genera Mycotrupes
and Peltotrupes (see the discussions under these ge-
nera). With a few rare exceptions, it appears too pre-
mature to base elaborate conclusions and speculations
on the inadequate, fragmentary data which we now
have. When we find what appears to be a disjunction
in the distribution of a species, we cannot determine
if this is a natural situation or if it is due to incom-
plete sampling. If we arbitrarily decide that this is a
natural pattern, then we can further speculate about
such things as limiting factors, geological history,
isolating mechanisms, etc. Even in Florida, where the

Scarabaeidae probably have been collected more in-
tensively than any other state, there are large areas
which have only barely been sampled. This is espe-
cially true for the western panhandle, although this
is a critical area and appears to be a zone of overlap
between several northern and southern populations.
States such as West Virginia and Wyoming have been
so poorly studied that nearly every distribution map
has a void of records from them. Considerable field
work will be necessary before the distributional pat-
terns can be used as if they reflected a natural
In the present work I have attempted to present
the distributional information available, pointing out
any factors (soil, climate, host association, sampling
error, etc.) which might be limiting. However, the
data are not sufficient to allow for many general-
Space does not permit a detailed discussion of
this interesting, but complicated, subject of biogeog-
raphy. Considerable work being conducted on other
groups of animals and, in paleontology, will undoubt-
edly produce new and critical data on the past and
present distributions of Florida animals. For addi-
tional information on speciation in the southeastern
Coastal Plains, the geological history of the area, and
the biogeography of Florida, the reader is referred to
the following papers: Alt and Brooks (1965); Berner
(1950); Byers (1930); Carr (1940); Carr and Goin
(1955); Clench and Turner (1956); Cooke (1939,
1945); Davis (1942, 1943, 1960); Gano (1917); Goin
(1958); Harper (1914, 1921, 1926, 1927); Henderson
(1939); Hobbs (1942); Howden (1963, 1966a); Hub-
bell (1932, 1954, 1961); Hubbell, Laessle, and Dick-
inson (1956); Kurz (1942); Laessle (1942, 1958);
Landsberg (1949); MacNeill (1951); McCrone (1963);
Mitchell (1963); Mount (1963, 1965); Neill (1957);
Safford (1919); Schuchert (1910, 1929, 1935); Schwarz
(1888); Vaughan (1910); West and Arnold (1956);
Young (1954).



Fig. 17. Average approximate hours of frost per year (from
Atlas of Florida).

Fig. 18. General soil distribution (after Atlas of Florida).
1) Northern upland types: well-drained loams and sands on
undulating uplands; forests of long-leaf pine, loblolly pine,
and hardwoods. 2) Northern slope types: well-drained to overly
drained sands and loamy sands; rolling, sloping, forested
land. 3) Central upland types: well-drained to overly drained
sands; rolling uplands, some sinkholes; slash pine and hard-
wood forests. 4) Central upland potash types: like 3 but
rich in potash. 5) Flatwood lowlands: poorly drained sands
or loamy sands; level land with many ponds and swamps;
mostly flatwoods or grass. 6) Southern limestone types:
poorly drained sands and loamy sands over limestone; flat-
woods or grass. 7) Swamp marsh and bottom lands; cypress,
gum, or marsh grass.

Nearly all of the drawings were made by the
author with the aid of a grid in the ocular of a stereo-
scopic microscope (10X and 15X oculars, .66, 1.3,
2.5, 4.0, and 5.0X objectives). The exceptions are Fig.
372-379 which are modified from published figures,
and they are appropriately acknowledged in the leg-
ends. The scale lines indicate length in relation to
the beetle size, regardless of reduction.
A general habitus drawing is presented for each
of the 32 genera treated in this volume. The repre-
sentatives were selected primarily because they are
typical for each genus, or, in some cases (e.g., Ataenius
saramari), because the species had not been illustrated
previously. If they serve their purpose, most Florida
genera of Laparosticti can be identified by a perusal
of these 32 drawings. They are made on a pebble-
surfaced scratchboard (Ross Board #2, C. J. Ross Co.,
Philadelphia) with India ink. The shading was done
with scalpels, and the fine lines and white setae were
made with a fine steel point set in a wooden handle.
Most of them are reduced from one-half to two-thirds
of the original.
Additional line drawings are provided for charac-
ters which clarify the keys or text. Near the comple-
tion of the manuscript I had access to a Cambridge
Instruments "Stereoscan" (SEM) through the courtesy
of the USDA (P. S. Callahan and Patricia Carlisle).
The illustrations from this source have enabled me to
clarify various taxonomic characters and to distinguish
some closely related species which could be conveyed
by no other means. The original photographs were
taken on a Polaroid camera attached to the SEM.
Various other photographs, especially of typical
habitats, are included. Most of these were taken by
the author, but the source for the others is acknowl-
edged in the legends.

Fig. 19. Average annual rainfall in inches (from Florida Wild-
life Magazine).


Although the family is one of the better known of
the larger families of beetles, many genera need re-
vising. For example, the large genera Aphodius and
Ataenius have not been revised, even for the U. S.,
since the time of Horn (1887) (although a revision
of Ataenius by Cartwright is in press). I have indicated
the status of the classification under each genus treated
here and suggested the need for generic revision where
it is apparent.
The higher categories (i.e., subfamilies, tribes, sub-
tribes, and genera) of the Scarabaeidae have rarely
been studied on a world basis, most authors limiting
their work to one geographic area. For this reason
it is very difficult to establish a good general scheme
of higher categories with any degree of phylogenetic
arrangement. The family is so large that it would be
an overwhelming task to study critically the entire
assemblage. Recently there have been some attempts
to establish a phylogenetic arrangement within certain
subfamilies (e.g., Scarabaeinae by Halffter and Mat-
thews, 1966).
Many European authors, including Crowson (1955),
have treated as families several of the groups recog-
nized here as subfamilies. Basically I have followed
the arrangement of families and subfamilies of the
superfamily Scarabaeoidea listed by Arnett (1962),
with the exception of recognizing the Trogidae as a
full family:

Subfamilies of
Families Scarabaeidae




Boving and Craighead (1931)
larval forms of Coleoptera,
Scarabaeoidea as follows:


Subfamilies of

,in their synopsis of the
treated the superfamily

Subfamilies of


Edwards (1949a), in treating the beetles east of the
Great Plains, arranged the Scarabaeoidea as follows:

The arrangement of the superfamily in several
major references is presented in the following discus-
sion. The most recent catalogue of the Coleoptera of
America north of Mexico is that of Leng (1920), with
five supplements (Leng and Mutchler, 1927, 1933;
Blackwelder, 1939; Blackwelder and Blackwelder,
1948). The superfamily Scarabaeoidea is arranged
as follows: *



Subfamilies of

In the "Coleopterorum Catalogus" (1910-37), a world
catalogue authored by various specialists, the Scara-
baeoidea are arranged as follows:


Crowson (1955), in his treatise on the natural classi-
fication of the families of Coleoptera, treated the
Scarabaeoidea as follows:

Subfamilies of
Families Scarabaeidae



In the present work, of a limited faunal region, no
attempt has been made to reevaluate the status of the
higher categories between subfamily and genus. In
most cases the latest, and presumably the most thor-
ough, arrangement is provisionally followed. Where
obvious inconsistencies have been noted they are
pointed out.
Ritcher (1969a) in an extensive survey of the
abdominal spiracles, suggested that the ". . . terms


Subfamilies of

Fig. 21. A view looking west from the bluffs on the east
side of the Apalachicola River at Torreya State Park. This
river appears to be the dividing line between several sub-
species of insects as well as a corridor for the southern ex-
tension of many northern elements.

Laparosticti and Pleurosticti should be abandoned."
He concluded that, within the Scarabaeoidea and even
within subfamilies, repeated evolutionary changes in
structure and number of functional spiracles do not
indicate the presence of two series. His evidence is
strong, and the two groups are probably not at all
natural. I had used the term Laparosticti in my origi-
nal manuscript primarily to avoid listing all the sub-
families each time they were to be mentioned. Un-
fortunately Ritcher proposed no new classification, and
therefore there is no other convenient term to use for
this group. I have maintained the use of the terms
here for convenience only, fully cognizant that they
may not reflect natural groupings.
The posterior procoxal bridge has been studied by
Ritchter (1969b) and used to interpret some higher
categories. He found that the Passalidae were so
distinctly different from other Scarabaeoidea that, along
with larval differences, this structure suggested that
they ". . . may have arisen from an earlier offshoot

Fig. 20. Habitat along the Tamiami Trail near Ochopee.
Much of southern Florida is similar to this area where few
scarabs occur except in the hammocks shown in the distance.

ARt A.113 -Cli

Fig. 22. Dune habitat at St. Andrews State Park, Fla., char-
acterized by scrub oak (Quercus virginiana maritima) and
rosemary (Ceratiola ericoides).

a Z u s
�AII -,

Fig. 23. Close up of a rosemary bush, showing the roots
exposed by wind and shifting sands of the active dunes. This
is a characteristic plant of many scrub habitats on fossil

of the ancestral scarabaeiform stock, not as a direct
offshoot of Lucanidae as suggested by Crowson." The
procoxal cavities are closed posteriorly in all Scara-
baeoidea studied, except in Pleocoma. He suggested
that the open procoxal cavities, as well as larval differ-
ences, ". . . indicates that Pleocoma is probably the
least specialized genus of the Scarabaeoidea."
In larvae, Hinton (1967) stated that a closing
apparatus for the spiracles is present in the Trogidae,
Lucanidae, Passalidae, and Glaphyrinae but absent in
the Geotrupidae and Scarabaeidae. He also stated that
the ecdysial process of the spiracles is of the elateroid
type in primitive Scarabaeoidea, whereas it has a bulla
that carries the ecdysial tube in specialized Scara-
baeoidea. For the latter he coined the term "pseudo-
panorpoid." Larval spiracles are crilfriform (Fig.

Fig. 24. General view of the habitat of Mycotrupes cart-
wrighti Ols. & Hub. Location: about six miles east of Talla-
hassee, Fla. in the "Red Hills" region. Specimens were col-
lected in bait traps placed in the roadway.

Fig. 25-26. Stereoscan photos of larval spiracles of Strategus
antaeus (Drury). 25) magnified 39X, 26) 83X.

Fig. 27-29. Stereoscan photos of an interior view of the
larval spiracle of Strategus antaeus (Drury) shown in Fig.
25-26. 27) note pattern of tracheal tubes, 81X, 28) 274X,
29) 713X.

25-29) in all scarabaeoid larvae except one group of
Trogidae. Baker (1968) revived the generic name
Omorgus for the Trox suberosus group of Vaurie (1955)
for the species of Trox having larvae with cribriform
spiracles, and those with biforous spiracles are retained
in Trox. Hinton (1967) stated that this biforous con-
dition is more primitive and resembles the elateroid
spiracles of other beetle larvae. Using this criterion
(larval spiracles) the genus Trox would then appear
to be the most primitive scarabaeoid type.
Baker (1968:2), while studying larvae of Trox,
stated that he believed this group was a subfamily of
the Scarabaeidae. This is the conservative approach,
and yet he proceeded to elevate one of the subgroups
of Trox to generic status. Unfortunately no complete
phylogeny is known for any insect, and the relative
positions of higher taxonomic categories will always
be merely subjective opinions.
Virkki (1957) discussed the evolutionary signifi-
cance of the testis follicle in the Scarabaeoidea (Fig.
30). He concluded that three main form groups were
recognized: 1) differences in follicle length (typical
Scarabaeinae), 2) follicles always spherical, but vary-
ing in size (typical representatives are Aphodiinae),
3) follicles differing also in relative breadth (higher
phytophagous scarabs).



r r



Fig. 30. Relationships of scarab groups according to follicle
structure (after Virkki, 1957). The representatives studied of
Melolonthinae were Hoplia, Melolontha, Amphimallon, and
Serica; Rutelinae were Anisoplia and Blitopertha; Cetoniinae
were Cetonia, Potosia, Oxythyrea, Tropinota, and Trichius.

In a study of scarab spermatogenesis Virkki (1966)
reported that Psammodius and Aegialia had similar
appearing karyotypes, but their extrachromosomal
cytology was quite different. Aegialia has conspicuous
diplocentric growth of the spermatozoa, suggesting " ...
a close relationship with Aphodius and Onthophagus,
and especially to Aphodius." He (1966:340) reported
that the scarab, Lichnanthe rathvoni (Lec.), has the
earliest spermatogenesis known to occur in beetles.
The process probably begins in the prepupa, because it
was completed and no gonia remained in nine-day-old

Generic names contribute to considerable nomencla-
tural confusion since they are a part of the binomial
system. I believe that genera do not exist in nature
(although species do) and are merely subjective cate-
gories that assist in organizing the species into group-
ings ("pigeonholing"). In some cases there are clear-
cut dividing lines between such groups, but more often
there is a gradual transition. Because of the subjec-
tive nature of the generic concept, many species have
been moved back and forth from one genus to the
next, creating numerous nomenclatural problems, es-
pecially homonymy. This is one of the factors which
led Michener (1964) to suggest the idea of "uninomial
No revolutionary procedures will be adopted here,
but the problem becomes a very real one when trying
to decide what generic name to use for some of the
Florida species. This is especially true in the subtribe
Canthonina, where the large genus Canthon recently
has been fragmented, particularly by Martinez, Pereira,
and Halffter. Other recent authors, such as Howden
and Matthews, have suggested that many of these new
genera represent only "species groups" and should not
be accorded generic status. As long as there are two
schools of thought on this subject, and these authors
are publishing contemporaneously, there will be no
real solution to the problem, and nomenclature will
suffer. This situation is discussed further under the
tribal treatment.
Many nomenclatural problems were encountered
during the course of this work, several of which could
not be resolved because of the unavailability of type
specimens or literature. Certain others will require sub-
mission to the International Commission on Zoological
Nomenclature for an official opinion. Good examples
of the several kinds of problems can be found under
the discussion of the genus Ochodaeus.

Like nearly all other aspects of the family Scara-
baeidae, the morphology is quite diverse and it is
difficult to make many generalizations. The family was
divided into two sections-the Laparosticti and Pleuro-
sticti-based on the position of the abdominal spiracles
and the relative positioning of the posterior legs, as
has been discussed earlier.
The family varies tremendously in size of individ-
uals, from 1.9mm (.075 inch) for Pleurophorus longulus
Cartwr. to nearly 15cm (6.0 inches) for Goliathus.
The latter is probably the largest insect known (based
on bulk). In Florida, the maximum size of 5.7 cm
(2.25 inches) is attained by Dynastes tityus (L.).
Although most of the species are dull colored, the
family contains many with beautiful metallic colors,
rivaling the showiness of the most colorful Lepidoptera.
Some scarabs even have scale-like setae on the elytra
similar to those of butterflies (Fig. 31). The general
body shape is also variable, from the globular
Acanthocerinae to the elongate Aphodiinae. Some idea
of the diversity, in the Laparosticti alone, can be
determined by a perusal of the habitus drawings of
the genera treated here.

Fig. 31-32. Stereoscan photos of scale-like setae on the
elytra of Polyphylla occidentalis (L.): 31) 85X, 32) 750X.

Beetles of this family have been popular with
collectors because of their bright colors and the great
development of horns in several groups (especially in
males of Dynastinae, Scarabaeinae, and Geotrupinae).
Although some of the horns are fantastically large and
bizarre, and have been the subject of much admiration
and speculation (Arrow, 1951), little information is
available on the functional significance of these
Beebe (1947), in a study of the largest horn-bear-
ing scarab in the New World, Dynastes hercules (L.),
found that the male horns were used in combat and
in transporting the opponent and, less frequently, the
female. He also studied minor and major males in
captivity and concluded that ".. . copulations of the
minor were as complete and successful as those of
the majors." There is therefore little to suggest that
those with maximum horn development have much evo-
lutionary (selective) advantage over those with minor

the tip of the abdomen than to the middle pair of legs.
As a consequence, the metathorax is often enlarged
and elongate. In the Pleurosticti the posterior legs are
situated near the middle of the body, sometimes very
close to the middle pair of legs. In this group the
metathorax is often broader and shortened.

Fig. 33. Stereoscan photos of right antenna of Polyphylla
occidentalis (L.). The lamellate antenna is typical of the Scara-
baeidae (Lamellicornia); the maximum of seven lamellae is
found in Polyphylla. Arrow indicates area enlarged in Fig.

horn development. On one occasion he found a male
successfully lifted and transported twice his own
weight. He concluded that encounters between male
hercules beetles were ". . . usually rather brief, and
are dependent on the willingness of each to fight.",
and that there was a ". . . complete absence of any
courtship or display by the male in respect to the
All species have a lamellate antennal club arranged
so that the lamellae can be expanded (Fig. 33) or
appressed (Fig. 46) to form a compact club. The
antennae, inserted in front of the eyes below the frons,
contain from seven to eleven segments, the first of
which is often enlarged, elongate, and covered with
elongate, stiff setae. The club is usually composed
of three segments (Fig. 45, 53, 54), but contains as
many as seven in the genus Polyphylla (Fig. 33, 46).
The club is variously shaped, from ball-like in Bolbo-
ceras (Fig. 54) to asymmetrical in Phanaeus (Fig. 45)
or somewhat cup-shaped in Hybosorus (Fig. 317-18).
The antennae are the principal olfactory receptors
(Fig. 33-35), although the maxillary palpi serve at
least as secondary receptors (Landin, 1961).
The legs, especially the anterior ones, of nearly all
species, are adapted for digging fossoriall) (Fig. 49,
52, 179-80, 354-55, 362-63). The tarsal formula for
the family is given as 5-5-5, but the anterior tarsi are
wanting in a few cases (e.g., Deltochilum and male
Phanaeus). The posterior legs of the Laparosticti are
situated behind the middle of the body, usually nearer




Fig. 34-35. Stereoscan photos of Polyphylla occidentalis (L.)
antenna. Enlargement of area shown at the arrow in Fig. 33;
34) 128X, 35) 257X. The numerous sensors probably per-
ceive various physical and chemical stimuli (e.g., odors, infra-
red and ultra-violet radiation, sound).

Fig. 36. Stereoscan photos of basal antenna segment of
Polyphylla occidentalis (L.) (133X). Arrow 1 indicates area
enlarged in Fig. 37; arrow 2 indicates area enlarged in Fig.


Fig. 37, Stereoscan photos of Polyphylla occidentalis (L.).
Base of sensor on basal antenna segment (enlargement of
arrow 1 in Fig. 36) (127X).

Fig. 38. Stereoscan photo of Polyphylla occidentalis (L.).
Middle area of sensor on basal antenna segment (enlarge-
ment of arrow 2 in Fig. 36) (2475X).

The anterior coxae are large, transverse, and some-
times prominent and conical. The anterior tibiae are
usually broad, flattened, and dentate on the outside.
The apex of the anterior tibia, on the inside, contains
a single spur which is often modified in the males.
The middle coxae are relatively large and transverse,
but not prominent. The posterior coxae are flat and
transverse. The middle and posterior tibiae are vari-
able in form; they are narrow, curved, and fitted for
ball rolling in the Scarabaeini (Fig. 51, 84); flattened
and sickle-shaped, and the surface covered with incised
lines in the Acanthocerinae (Fig. 392-95, 400-401);
in all other groups they are generally expanded at the
apex or nearly straight and parallel. The posterior
tibiae have two apical spurs except in the Scarabaeini
(where only the genus Melanocanthon has two). The
tarsal claws vary from simple to toothed (Fig. 47-48).
Although there is usually a bisetose onychium, it is
wanting in the Acanthocerinae, Aphodiinae, and some
Matthews (1972) discussed the interesting modifi-
cations of tarsal claws of the group of Onthophagus
previously known as Macropocopris. He found that the
prehensile claws (used for holding on to hairs of
wallabies, kangaroos, and goats) were derived in two
ways: 1) the last tarsal segment bears a double row
of stiff bristles ventrally which act in apposition to the
claws; pulvillus present; 2) the last tarsal segment
bears a spur ventrally which acts in apposition to the
claws; pulvillus absent or greatly reduced.
The mouthparts are highly variable in form, from
the nearly membranous type of the Scarabaeinae and




Fig. 39-42. Key characters for subfamilies, tribes, and ge-
nera: 39) Ventral view of the abdomen of Cloeotus apho-
dioides (III.). Note the five visible segments. 40) Ventral
view of the abdomen of Geotrupes egeriel Germ. Note the
six visible segments. 41) Dorsal view of right posterior leg
of Aphodius fimetarius (L.). 42) Dorsal view of the right
posterior leg of Ataenius alternatus (Melsh.). Note the ab-
sence of scalloping and transverse carinae.

Aphodiinae, to the hard, well-developed mandibles of
the Geotrupinae and Dynastinae. The mouthparts are
described in detail by Hardenberg (1907) from ex-
amples throughout the family. Landin (1961) dis-
cussed these organs in the Aphodiinae, and Miller
(1961), and Halffter and Matthews (1966) considered
these structures in several Scarabaeinae. The maxillary
palpi are slender, four-segmented, with the apical seg-
ment the largest. They often contain odor and, prob-
ably, taste receptors. The labial palpi are three-seg-
mented. Halffter and Matthews (1966:243) stated
that ". . . the mouthparts [of Scarabaeinae] as a whole
must have an extraordinary ability to taste, sort, and
screen very small particles down to the size of bac-
teria." Miller (1961) found the gut to contain particles
in suspension, varying in diameter from two microns
(Onthophagus) to 16 microns (Dichotomius).
There has been little work done on the internal
anatomy of the dung beetles, except for the few papers
that are summarized by Halffter and Matthews (1966).

Fig. 43-49. Key characters for subfamilies, tribes, and ge-
nera: 43) Lateral body outline of Cloeotus globosus (Say). Note
the compact, globular shape characteristic of the Acanthoceri-
nae. 44) Lateral body outline of Ataenius alternates (Melsh.).
Note the elongate, cylindrical shape of the Aphodiinae. 45)
Dorsal view of the right antenna of a male Phanaeus igneus
floridanus d'Ols. Note the three-segmented club. 46) Dorsal
view of the right antenna of a male Polyphylla pubescens
Cartwr. Note the seven-segmented club. 47) Ventral view of
the right posterior tarsus of Phanaeus igneus floridanus d'Ols.
Note the absence of terminal claws. 48) Lateral view of the
tip of the left posterior tarsus of Phyllophaga latifrons (Lec.).
Note the cleft tarsal claws. 49) Dorsal view of the right
anterior tibia of a male Phanaeus igneus (loridanus d'Ols.
Note the absence of the tarsus.

50 52

53 51
53 545

55 56 57 58

Fig. 50-58. Key characters for subfamilies, tribes, and gen-
era: 50) Dorsal view of right posterior tibia of Dichotomius
carolinus (L.). Note the expanded apex. 51) Dorsal view of
right posterior tibia of a male Deltochilum gibbosum (Fab.).
Note the long, slender form, and the apex is not noticeably
expanded. 52) Dorsal view of anterior tibia of a female
Deltochilum gibbosum (Fab.). Note the absence of a tarsus.
53) Dorsal view of right antenna of Geotrupes egeriei Germ.
54) Dorsal view of right antenna of Bolboceras floridensis
(Wallis). 55) Dorsal view of right posterior tibial apex and
tarsus of Pleurophorus longulus Cartwr. 56) Dorsal view of
right posterior tibial apex and tarsus of Psammodius malkini
Cartwr. 57) Dorsal view of right posterior tibia and tarsus
of Melanocanthon granulifer Schmidt. 58) Dorsal view of right
posterior tibia and tarsus of Boreocanthon depressipennis

These authors discuss the morphological adaptations
for coprophagy in the adults and larvae of the Scara-
baeinae. Becton (1930) described the alimentary tract
of Phanaeus vindex MacL., and Miller (1961) de-
scribed the same structure in Canthon pilularius (L.).
The female reproductive system has been examined
only for a few representative genera. It is similar to
that of other Coleoptera in general, but in all Scara-
baeinae examined, there is only one ovary, and it is
reduced to a single ovariole on the left side. Robertson
(1961) indicated that this is the only group of Coleop-
tera showing such a degree of ovarian reduction.
The ma!e genitalia are useful for distinguishing
species of certain genera (e.g., Melanocanthon, Bolbo-
cerosoma, and Phyllophaga). However, there has been
no extensive comparative study within the family.

There are a few studies comparing these organs be-
tween families of Coleoptera (Sharp and Muir, 1912;
Snodgrass, 1957).
Although the female genitalia probably are diag-
nostic, they have been studied only in a few cases
(e.g., Phyllophaga). Tanner (1927) discussed these
organs throughout the Coleoptera, but few examples of
Scarabaeidae were studied. I have been interested in
this subject for some time, having accumulated several
hundred slides for future studies. It is premature to
make many broad generalizations, but it appears that
a great many species can be separated easily on the
basis of the female genitalia.
There are no comparative morphological studies on
the family as a whole. The subfamilies Scarabaeinae,
Aphodiinae, and Geotrupinae were studied compara-
tively by Mohr (1930), but using only a single species
from each. Much more is needed in the way of com-
parative morphology before there can be a logical
treatment of the higher categories within the family.
Scarab larvae are usually C-shaped white grubs
(Fig. 59), with the head capsule fairly large, hard, and
dark colored (Fig. 61). However, many of the Scara-

ian e .Cm.

Fig. 60. Larval raster of the last ventral abdominal segment,
a character useful in distinguishing larvae, of many species.
Phyllophaga bruneri Chapin.


Fig. 61. Typical scarab larval head, showing morphological
structures. Phyllophaga bruneri Chapin.

Fig. 59. Typical C-shaped scarab larva. Lateral view of third
instar of Phyllophaga bruneri Chapin setaee omitted).

baeinae and Geotrupinae have a "hump-backed" ap-
pearance (Fig. 10). It is not within the scope of the
present study to discuss the immature stages in detail.
However, I have indicated under each species treatment
whether they are known and, if so, I have briefly
characterized them. The most useful taxonomic char-
acters are found on the mouthparts (especially the
epipharynx) (Fig. 62, 64) and the venter of the last
abdominal segment (raster) (Fig. 60).' During the
latter part of this study an excellent book appeared,


Fig. 62. Larval epipharynx, showing taxonomic structures
used to distinguish species. Phyllophaga bruneri Chapin.

summarizing the described North American scarab
larvae (Ritcher, 1966).
Puchkova (1966) reported pupal structures in Sca-
rabaeidae which apparently correspond to rudimentary
tergopleural glands and others corresponding to an
external ovipositor. He also discovered structures on
many scarab pupal abdomens that help clarify the
morphogenesis of the pygidium. He suggested that the
shape of the pupa indicates that scarab ancestors had
campodeiform larvae, and that their present C-shape
arose secondarily in connection with an increase of
larval body mass.


Fig. 63. Typical scarab pupa. Ventral view of Phyllophaga
bruneri Chapin.

The area encompassed by this subheading includes
ecology and ethology, but they are so interwoven as to
be nearly impossible to discuss separately. Ecology
in its broadest sense includes all aspects of the habits
and behavior of an organism in relation to both its
abiotic and biotic environment. An attempt is made
to summarize our knowledge of this subject under the
individual species treatments, and only brief general-
izations will be made here.
The family, as mentioned earlier, can be divided
on the basis of food habits into two major groups
which once were thought to coincide with morphologi-
cal divisions. The Pleurosticti feed primarily on live
plant materials as adults, and the larvae usually feed
on plant roots or rotting wood. As,they are not to be
treated in this work, they will not be discussed further

here. The Laparosticti, on the other hand, feed pri-
marily on dung or decaying plant and animal materials
as adults, and in many cases larval food of the same
materials is provisioned by the adults.
Matthews (1963), in referring to the common
Canthon pilularius (L.), stated "The autecology of this
species has not been properly investigated (nor has
that of any other American coprophage)." The same
statement could apply to nearly all members of the
family, with the possible exception of a few economi-
cally important species such as the Japanese beetle
(Popillia japonica Newm.).
Even though we are far from knowing the details
about the ecology of most scarabs, the literature on the
subject is extensive. Ritcher (1958) briefly reviewed
174 papers on the "Biology of Scarabaeidae." Landin
(1961 & 1968) published the excellent books on the
ecology of the subfamily Aphodiinae, but these works
involved only European species. However, much of
the general information on ecology of the dung-feeding
species is applicable to Florida species. The most
recent and most significant publication on the subject
is the volume by Halffter and Matthews (1966) on
"The Natural History of the Dung Beetles of the Sub-
family Scarabaeinae." This work provides an exhaus-
tive review of the literature of the group, as well as
much valuable original data. Howden, in a series of
papers (especially 1955a & 1964), has done an out-
standing job of summarizing our knowledge of the
North American Geotrupinae. The remaining three
subfamilies of Laparosticti treated here (Ochodaeinae,
Hybosorinae, Acanthocerinae) are very poorly known
biologically. The food habits of both adults and larvae
of the Florida species are unknown.

Fig. 64. Stereoscan photo of larval epipharynx of Strategus
antaeus (Drury) (43X).

1) Temperature is limiting in two directions. High
temperatures cause immobilization and death; low
temperatures cause inactivity and cold stupor; the
average optimum is between +14 degrees and
-17 degrees C., with a wide tolerance range.
2) Relative humidity is limiting in one direction.
Low humidity causes decreasing flight activity, and
dessication can be fatal. High humidity favors
flight activity, with the optimum between 70 and
A 100% RH.
3) Light (solar radiation) does not seem to have
a limiting effect. They seem to "prefer" darkness,
but are able to adapt from flight in darkness to
daytime flight.
4) An endogenous system ("internal clock") ap-
pears to maintain some control, especially under
optimum conditions.
5) The optimum climatic factors for flight, reflected
by response from the endogenous rhythm, are: a)
moderate temperature, b) high relative humidity, c)
and darkness.
6) The "wide ecological amplitude" of most
Aphodius ". . . considerably adds to the ability of
the species to disperse, often over wide areas, and
even to settle down in new, often far distant con-

In Florida there are few species for which any
details of the ecology or ethology are known. There
are some interesting close associations between some

Gophers polyphemus Doudin

Fig. 65-66. Stereoscan photos of stridulatory area on left
mandible of Strategus antaeus (Drury): 65) 45X, 66) 190X.

Landin (1968) has published the only extensive 67
study of this aspect of behavior, but he treated only
the genus Aphodius in Sweden. He reached the follow- ,,004
ing conclusions based on field studies and laboratory Fig. 67-68. Distribution of the gopher tortoise (based on data
experiments: supplied by W. Auffenberg).

One species (Ataenius new species near brevinotus
Chapin) has been found in fox squirrel (Sciurus niger
L.) nests. One species (Ataenius insculptus Horn)
has been found only in deer droppings, and another
(Aphotaenius carolinus (Van D.)) is found primarily
in deer droppings. Two species [Euparia castanea Serv.
and Myrmecaphodius excavaticollis (Blanchard)], are
myrmecophilous, and they occur respectively in the
nests of the fire ants Solenopsis geminata (Fab.) and
Solenopsis invicta Buren (Fig. 76-77).

In many of the Scarabaeinae and Geotrupinae the
adults provision the larvae with food in a subterranean
chamber (nidification) (Fig. 9-11). This behavior is
carried to the ultimate in some species of Copris, in
-� t2( ..mU"

Fig. 69. Habitat of packrats (Neotoma floridana floridana)
in a small cave in Florida Caverns State Park. Scarabs found
in packrat droppings here were Onthophagus orpheus Panz.
and Aphotaenius carolinus (Van D.).

Fig. 71. Stick nest of packrats (Neotoma floridana small)
on North Key Largo, Fla. Onthophagus orpheus Panz. and
Ataenius brevicollis Wallaston were found in this nest.

Fig. 70. Close up of packrat droppings on a rock at the
habitat shown in Fig. 69, in which the two scarabs men-
tioned above were found. (Both photos by F. W. Mead)
Scarabaeinae and Aphodiinae and other animals which
should provide valuable information once they are
more thoroughly understood. There are three species
(Onthophagus polyphemi Hubbard, Copris gopheri
Hubbard, and Aphodius troglodytes Hubbard) found
only with the gopher tortoise (Gopherus polyphemus
(Daudin) (Fig. 67-68). Three species (Aphodius
laevigatus Hald., Aphodius aegrotus Horn, and Apho-
dius haldemani Horn) are found only with pocket
gophers (Geomys spp.) (Fig. 74-75, distribution map
based on unpublished data provided by C. H. Handley
and E. V. Komarek). Three species (Onthophagus
orpheus (Panz.), Pseudocanthon perplexus (Lec.), and
Ataenius brevicollis Wollaston) have been found in
packrat (Neotoma spp.) nests (Fig. ,69-73), although
only the latter seems to be confined to such habitats.

Fig. 72. Packrat nest shown in Fig. 71, after removal of the
stick covering. Most of the scarabs were found in droppings
in the cavity at the base of the tree.

Distribution of pack rats
[Neotoma floridana]
in Florida

E1 N. f. floridana
N. f. illinoensis
N. f. small

packrats in Florida (modified from

Fig. 74. Mounds of the pocket gopher (Geomys pinetis
austrinus), in the burrows of which are found Aphodius
aegrotus Horn (= geomysi), Aphodius laevigatus Hald., and
Aphodius haldemani Horn. Location: about 10 miles north of
Ocala, Fla.

which the adult female stays with the brood ball
through pupation. The dung mass is continually
smoothed on the outside, preventing the growth of
mold. Halffter and Matthews (1966) discussed nidifi-
cation in the Scarabaeinae, and Howden (1955a) re-
viewed this behavior in the Geotrupinae.

Many of the Laparosticti are subsocial or colonial,
especially the Geotrupinae. Most of the dung-feeding
species are often found together in a single pile of
cow dung. A great many of the species stridulate
(Fig. 78), but the behavioral significance of sound
production has not been thoroughly investigated. Arrow

Fig. 75. Distribution of Florida pocket gophers (based on data
supplied by C. H. Handle and E. V. Komarek). 1) Geomys
colonus Bangs, 2) Geomys cumberlandius Bangs, 3) Geomys
pinetis pinetis Rafinesque, 4) Geomys pinetis floridanus (Aud.
& Bach.), 5) Geomys pinetis mobilensis Merriam, 6) Geomys
pinetis austrinus Bangs, 7) Geomys pinetis goffi Sherman.

(1904) described the sound producing organs for a
number of species in the family, and the evolutionary
significance of sound production of beetles was dis-
cussed by Alexander, Moore, and Woodruff (1963).
This area of investigation offers a fertile field for
future research.

Anyone who has spent a few hours in a pasture,
witnessing the speed at which a cow dropping is
colonized by dung beetles, cannot help wondering about
the sense of smell in these beetles. Mohr (1943) re-
ported 1,097 specimens of Aphodius distinctus Mull.
in a single cow dropping within 2 hours. Lindquist
(1935) found 1,113 specimens of Aphodius lividus
(Oliv.) in a single cow dropping. I have collected as
many as 12 species of scarabs in a single cow drop-
ping, within 2 hours after deposition.
In Finland, Rainio (1966) found that there were
species of dung beetles which preferred each of the
following kinds of dung: horse, human, cow, and sheep.
However, he found that none were wholly specialized
to one type. Successionally he found that the numbers
of beetles reached a peak in: 1) horse dung after one
day, 2) sheep and pig dung after two days, and 3) cow
dung after three days. Although certain species pre-
ferred shaded habitats, he found that most species were
more abundant in exposed situations.
Although the perception of, and orientation to,
odors is one of the most important aspects of dung
beetle behavior, it has received very little attention.
For the Scarabaeinae, Halffter and Matthews (1966)
have stated, "Such aspects as the pattern of search

Fig. 73. Distribution of
Hall and Kelton).

It is likely that the initial odor stimulation may be
at a great distance from the source, but orientation is
confined to short distances. There are probably con-
siderable specific differences in the ability to perceive
and orient toward the food source. In general, stimula-
tion is accompanied by an "alarm" reaction, the head
being raised, the antennae extended, and the lamellae
of the club being spread apart. Flight is usually toward
the wind, and initially the track twists and turns. Once
a sufficient odor gradient is perceived, the track is
nearly a straight line to the source. We have very
little information on this aspect of behavior, but the
straight part of the track has been reported as 10
meters for Scarabaeus and 50 cm for Geotrupes (Fraen-
kel and Gunn, 1961:279), and 40 cm for Aphodius
(Landin, 1961:182). Some species land within short
distances and walk the rest of the way, while others
land directly on the dung. Although the details are
mostly unknown, there can be no doubt about their
success and speed of finding a food supply of dung.
Fig. 76. A pasture containing numerous mounds of the im- It would be especially interesting to learn how those
ported fire ant (Solenopsis invicta Buren). Location: Lowndes species associated with certain mammals and ants find
County, Alabama. (Photo by F. W. Mead) their hosts.
Dung beetles are interesting subjects for ecological
studies for several reasons: 1) dung is a rapidly chang-
ing microcommunity which forms a suitable subject for
succession; 2) dung, along with decaying plants and
animals, is one of the most universal habitats, being
found under a wide range of environmental conditions;
3) dung beetles have a wide geographical range and
occur nearly worldwide except for Antarctica and a
few oceanic islands; 4) they often occur in large num-
bers, making the study of populations feasible; 5)
many species have evolved unique commensal relation-
ships with other insects and mammals.

Several dung beetles have been introduced into
Australia from Africa to: 1) free pastures from ac-
cumulated dung, 2) fertilize the soil, and 3) control
pests. The idea of improving pastures with introduced
species was proposed by Bornemissza (1960), and his
Fig. 77. A cross section of an imported fire ant nest. Myrme- work continues at present. During the International
caphodius excavaticollis (Blanch.) is found in such nests. Congress of Entomology at Canberra, Australia (Au-
Euparia castanea Serv. is found in similar nests of the re- gust, 1972) much emphasis was placed on this work,
lated ant, Solenopsis geminata (Fab.). Photo by F. W. Mead) and an excellent educational film entitled "Dung down

flight, territory covered by a single beetle, altitude of ,
flight, distances at which the smell of food is first
perceived, etc., are nearly unknown." The same state-
ment is equally applicable to the dung-feeding species
of Aphodiinae and Geotrupinae.
The olfactory organs have been studied only in a
few cases (Warnke, 1934). It has been determined
by experimentation that the principal seat of olfaction 7,
is found in the antennal club, and to a much lesser -
degree in the maxillary and labial palpi. The two gen-
eral types of receptors consist of small conical cells '
which penetrate the cuticula and end externally as
pegs (sensilla basiconica), or as short setae (sensilla
trichodea). Another type, in the form of conical pegs Fig. 78. Audiospectrograph tracings of scarab sounds (dis-
arising from small pits (sensilla' coeloconicum), is turbance at 80-85' F.): a) Geotrupes blackburnii (Fab.), b)
found on the ultimate segment of the maxillary palpi. Cloeotus aphodioides (I11.).

under" was shown. The Commonwealth Scientific and
Industrial Research Organization (CSIRO) has spent
much time and effort on this project and maintains
a laboratory in South Africa. Here Bornemissza is
searching for species with the proper ecological re-
quirements to fill specific niches in Australia.
Woodruff (1972) summarized much of the CSIRO
work. In dry climates, dung often remains on the
surface for months. It is estimated that 200 million
dung pats are deposited daily by Australian cattle,
annually covering hundreds of thousands of acres of
potential pasture. Beetles can bury the dung within
48 hours and thus increase soil fertility by incorporat-
ing it into the soil. The beetles' mechanical disturb-
ance of the dung prevents completion of development
of several pest flies. The burial of dung reduces the
number of parasitic worm eggs and the resultant in-
fective larvae which reach grass blades on which they
might be ingested by cattle.
The most successful introduced species is Onthopha-
gus gazella (Fab.), a species earlier introduced into Ha-
waii to assist in hornfly control. In Australia it had col-
onized 400 kilometers in 2 years and appears to be
rapidly expanding its range. In the first year it
crossed 7 miles of water to colonize an island, and the
following year it reached an island 30 miles away.
Mass releases are still being made under carefully
controlled conditions. Eggs are collected from dung
balls in Africa, surface sterilized with a 3% formalde-
hyde solution, air freighted to Australia in special
containers, washed on arrival, and placed in hand-made
dung balls. The adults of this generation are used for
egg production only, and adults of the second genera-
tion are used for the mass releases.
Colonies of some African species (e.g., Onthophagus
gazella) are maintained in the USDA laboratory at
Kerrville, Texas, and specimens very recently were re-
leased in Texas (Drummond, personal communication).
The only introduced (accidental) dung beetle in Flor-
ida is Onthophagus depressus Har. Although it is well-
established, it appears to be of little consequence in
the pasture ecosystem (see discussion under this species
in the text).
Gillard (1967) has also discussed the role of copro-
phagous beetles in the pasture ecosystem. He pointed
out that most native, tropical, and subtropical pastures
have a relatively low nitrogen content compared with
improved temperate pastures. He found that unburied
dung dried by the sun had an 80% loss of nitrogen,
while that which is buried loses 5 to 15% by volatil-
ization. Although there are many complicated factors
to measure and assess, there is little doubt that the
dung beetle fauna contributes to conservation of nitro-
gen, resulting in increased soil fertility.

In general the dung beetles are seldom attacked
by parasitic Hymenoptera or Diptera. They often har-
bor mites of many kinds, but this association is
thought to be primarily phoretic. Halffter and Mat-
thews (1966:171-176) presented a list of the mites
associated with various species of Scarabaeinae. In

Fig. 79. A burrowing owl (Speotyto cunicularia floridana) al
the entrance to its burrow. Location: Sun Springs, Gilchrist
County, Fla. (Photo by C. T. Collins)

Fig. 80. Burrowing owl pellets containing insect remains. A
high percentage of the owl's diet is composed of dung
beetles, especially Dichotomius, Phanaeus, Canthon, Copris,
Peltotrupes, and Mycotrupes. (Photo by E. M. Collins, Jr.)

many cases the mites are thought to be predators on
fly larvae (Axtell, 1963) or nematodes (Stewart and
Davis, 1967). They represent several families, chief
of which are the Macrochelidae. These are actively
being studied by several workers, among whom are
Costa and Krantz.
Costa (1969) reported 183 species of gamasine
mites from Scarabaeidae, the greatest number from any
group of insect hosts. Among these were representa-
tives of the following six mite families (the number
of species indicated in parentheses): Dermanyssidae
(17), Eviphiidae (36), Macrochelidae (118), Pachy-
laelapidae (19), Parasitidae (8), and Rhodocaridae
(3). He found that various stages of the mites were
found on the beetle hosts. All stages were found in
Otopheidomenidae, deutonymphs only in Parasitidae

and Digamasellus spp., and female mites only in
Macrocheles spp., Dinogamasus spp. and Coleolaelaps
spp. In the genus Coleolaelaps (on phytophagous
scarabs), female mites are usually carried on the beetle,
whereas male and immature mites are found on the
beetle larvae (white grubs) underground. He recorded
19 species of mesostigmatic mites from a single species
of dung beetle [Copris hispanus (L.)], of which 3
appear to be host specific.
Krantz and Mellott (1968) studied an interesting
group of parasitic mites of the family Macrochelidae
which I found on two genera of burrowing beetles
(Peltotrupes and Mycotrupes) in Florida. They de-
scribed one new species from each genus of beetle
with the following statement regarding their relation-
ships: "There can be little doubt that one of the two
macrochelids to be described below evolved from the
other or, less likely, that both evolved from a com-
mon ancestor." These beetles are considered relictual
(Mycotrupes has lost the hind wings and contains dis-
junct populations), and a study of the beetle-mite
associations should produce interesting evolutionary
information on both groups.
Several animals are predaceous on scarabs, the most
notable being the Surinam toad (Bufo marinus L.)
which is successful in controlling May beetles (Phyl-
lophaga spp.) in Puerto Rico (Wolcott, 1937). There
is little doubt that other toads and frogs consume
their share of scarabs in Florida. In fact, one of the
few records of the rare Acanthocerus aeneus MacL. is
of a specimen from the stomach of a frog (Rana
sphenocephala Cope). Two skinks (Eumeces egregius
(Baird) and Neoseps reynoldsi Stejn.) were frequently
found in "push-ups" of Mycotrupes and Peltotrupes, but
they are not known to feed on the beetles (Mount,
1963). Animals such as skunks, opossums, and arma-
dillos undoubtedly feed on some scarab larvae and
adults, but little information is available on the quan-
tities involved. One of the most important predators
on dung beetles, in the limited areas where it occurs,
is the burrowing owl (Speotyto cunicularia floridana
Ridgway). Their pellets (Fig. 79-80) contain abun-
dant remains of Dichotomius, Phanaeus, Canthon,
Copris, Peltotrupes and Mycotrupes. During this study,
C. T. Collins and W. Courser supplied numerous owl
pellets, the data from which will form the basis of
a joint paper on the food habits of this owl in Florida.

Two things are immediately apparent when one
begins to analyze the distributions of Florida Laparo-
sticti: 1) the high percentage of species endemic to
the state, and 2) the paucity of overlap in both genera
and species with the West Indian fauna.
Of the 115 species and subspecies of Laparosticti
recorded from Florida, there are 24, or slightly more
than 20%, not recorded outside the state. It is likely
that some of these will eventually be found in south-
ern Georgia or Alabama, but they probably have a
fairly narrow range. There are 10 additional Florida
species that barely range outside the state. The en-
demics are probably the result of many factors, in-

eluding insular isolation during past geological periods
and unique environmental conditions (e.g., subtropical
climate). Only one genus of Laparosticti (Peltotrupes)
is entirely endemic to Florida.
Only 16 species (including the questionable Florida
record of Oniticellus cubiensis Lap.) are common to
Florida and the West Indies. All of these, except the
questionable species just mentioned and Hybosorus
illigeri Reiche, are in the subfamily Aphodiinae, and
most of them have a fairly wide distribution. One of
these species was thought to be Ataenius brevinotus
Chapin. Cartwright (in litt.) now believes that the
Florida specimens represent an undescribed species.
Another species, Ataenius luteomargo Chapin, known
from 11 islands of the West Indies, was recorded in
my dissertation for the first time from Key West,
Florida, where it was thought to be a recent introduc-
tion. However, Cartwright (in litt.) has now indicated
that these specimens represent a Cuban species,
Ataenius waltherhorni Balth., previously recorded from
the U.S. only from a questionable specimen from
"Everglade." Psammodius bidens Horn has been re-
corded from several areas from New Jersey to Florida,
and there is a single specimen known from Puerto
Rico. Hybosorus illigeri Reiche was recorded from
Jamaica by Howden (1970).
Of the 32 genera of Laparosticti known from
Florida, only 12 have representatives in the West
Indies. The subfamilies Geotrupinae and Ochodaeinae
have no genera or species common to both areas.
Ten of the Florida species, representing seven
genera, presumably have been introduced. One of
these (Onthophagus depressus Har.) has also been
found in Georgia and is from Africa. It was recently
recorded (Matthews, 1972) from Australia. Interest-
ingly, a related African species has been introduced
into Martinique in the Lesser Antilles (Matthews, 1966:
25). Oniticellus cubiensis Lap. occurs in the Bahamas,
Cuba, and Jamaica, and if the single Key West,
Florida, record (Blatchely, 1928:13) is valid, it was
probably introduced there. Another species (Myrmeca-
phodius excavaticollis (Blanch.)), here definitely re-
corded from the U.S. for the first time, was described
from Argentina in the nests of the fire ant Solenopsis
saevissima Fr. Smith). This ant and a similar, recently
described species (S. invicta Buren) were introduced
into the U. S. probably after 1915, and presumably
the beetle was introduced with its ant host. Two other
species (Ataenius simulator Har. and Psammodius
cruentus Har.)were described from South America and
are presumed to be fairly recent introductions to the U. S.
Hybosorus illigeri Reiche is widespread in the Old
World, from southern Europe to Africa, and it was
probably introduced into the U. S. at a very early
date (before 1848, when LeConte described it as
H. carolinus). The remaining four species are in the
genus Aphodius [fimetarius (L.), granarius (L.),
haemorrhoidalis (L.), and lividus (Oliv.)]. These are
all presumably European in origin, although they now
have wide ranges, probably as a result of distribution
by commerce. The many introductions of Aphodiinae
around the world have been discussed by Lindroth
(1957) and others.

For further details on distribution and zoogeogra-
phy, the reader should consult the discussions under
each species in the systematic account which follows.

There is a single report (Wickham, 1919) of a
fossil scarab from Florida. This is based on an elytral
fragment and a smaller fragment of Copris inemargin-
atus Blatchley from a Pleistocene deposit at Vero,
Florida. There has been considerable discussion of
the nature and age of these deposits (Sellards, 1916;
Weigel, 1962; Young, 1959). Until further specimens
are discovered in the undistributed portions of Stratum
2 (Weigel, 1962), I view this single record with scepti-
cism. None of the insects from this deposit are replaced
by minerals, and all appear as if they had recently
died. Only one species, a grouse locust, is not known
to occur in the vicinity of Vero today.
Many of the early collections from this site (in-
cluding the fragments of Copris inemarginatus) were
". .. carried out without stratigraphic control . . .
and "Instead, bones and artifacts were picked from
the sides of the canal banks." (Weigel, 1962:12). In
addition, animals such as pocket gophers, tortoises,
and even the beetles themselves can readily dig through
deposits without causing noticeable disturbance in the
beds. It is quite likely that at least some of the insects
were deposited in this manner. Regardless of the evi-
dence for other insects at this site, it is unwarranted at
this time to attach any great significance to the two
fragments of Copris inemarginatus. It is certainly too
premature to suggest that ". . . perhaps Copris inemar-
ginatus, a sort of living fossil, did once utilize the
dung of Equus, Mammut, and Mammuthus which are
also found as fossils with it at Vero." (Young,

The family Scarabaeidae contains some of the
most destructive beetles known, including the Japanese
beetle, Asiatic garden beetle, and European chafer.
However, nearly all of these are in the Pleurosticti
which well be treated later. The Laparosticti, with very
few exceptions, are believed to be either beneficial or
of little consequence to man.
Since the groups treated here are primarily dung-
feeders or scavengers, they are very useful in ridding
the countryside of waste materials. We often are not
aware of the numbers of species and individuals con-
stantly at work at this tremendous task. Hingston
(1923) remarked about the role played by this group
in Hindustan:

Were it not for Nature's scavengers the
East would be the cesspit of the world. Man
assuredly would annihilate himself in the
emanations from his own filth . . . But in-
comparably the chief of this great array
are the members of the enormous family of
Dung-beetles, the Scarabaeidae. They seek
the excrement of men and cattle, gather it

into nodules or rounded pellets, and bury it
beneath the surface of the soil. Since the
greater part of their life is spent hidden in
the earth or lodged in the substance of some
faecal mass, they are not obvious to every
eye. Nevertheless, they exist in prodigious
numbers, and the quantity of refuse which
they remove is immense, almost beyond be-
lief. So far as I can estimate by rough
observation, I believe that in certain active
seasons of the year two-thirds of the excre-
ment of this vast country must be carried by
these scarabs into the substance of the soil.
Without their valuable aid the land would
be an open sewer. Remember that it sup-
ports a teeming population of some 300
million souls. And, save for the few collected
in the cities, the whole of this great multi-
tude of people depends on the work of
Nature's scavengers to clear its filth away.
I will not enter into numerical details, but,
taking into account human ordure alone, I
believe, that in India during May and June
as much as forty or fifty thousand tons of
excrement must be carried by scarabs each
day into the soil. And this does not include
the dung of animals, which may easily
double or treble the amount. It seems an
almost incredible number. I advance it with
no claims to the strictest accuracy, but I
think it gives us a just impression of the
incalculable value of this tribe of beetles as
the leading scavenging army of the East.
Although little research has been conducted on the
benefits accrued by the incorporation of dung into the
soil, there can be no doubt about the increased aera-
tion and fertility as a result of this behavior. Lindquist
(1933) measured the amount of dung buried by cer-
tain species, and Bornemissza (1960) suggested that
dung beetles could improve pastures in Australia. (For
further discussion of this aspect of dung beetles see the
earlier section on Biology).
One of the beneficial aspects of dung beetles is a
result of the speed with which they can locate and
disrupt the dung. Larvae of the horn fly (Haematobia
irritans (L.) can survive only in undisturbed cow dung.
The feeding and tunneling of many scarabs often render
the dung unsuitable for complete development of the
fly larvae. There have been several attempts to intro-
duce dung beetles for this purpose into areas where
they were not abundant (e.g., Hawaii and Puerto Rico).
However, there have been no attempts to thoroughly
evaluate the results.
Many Laparosticti serve as intermediate hosts for
a variety of helminths (primarily Spiruroidea). Halffter
and Matthews (1966) summarized the literature on
this aspect of the Scarabaeinae. Some important animal
parasites which have one or more scarab intermediate
hosts are: Raillietina cesticillus (Molin) and Hymen-
olepis carioca (Magalhaes), intestinal parasites of
chickens and certain wild galliform birds; Ascarops
strongylina (Rudolphi), a stomach parasite of domestic
and wild swine; Spirura rytipleurites (Deslongchamps),
a stomach parasite of cat, dog, fox, skunk, and hedge-
hog; Spirocerca lupi (Rudolphi), a parasite of dog,

wolf, jackal, and fox; Physocephalus sexalatus (Molin)
and Gongylonema pulchrum Molin, parasites of a
variety of wild and domestic animals; Macracanthor-
hynchus hirudinaceus (Pallas), an intestinal parasite
of swine, carnivores, monkeys, and in one case, man.
Miller (1954, 1961) studied the dung beetles found
on human feces in Georgia and concluded that
". . . . dung beetles may constitute an important factor
in the epidemiology of hookworm and other enteric
parasites of man." Stewart and Davis (1967) discussed
the "consortisms" which exist between swine, nema-
todes, dung beetles, and mites. It appears that there
are many complicated close associations of this sort
which will require detailed studies before any conclu-
sions can be reached.
Although dung beetles serve as intermediate hosts
for some vertebrate parasites, they are also involved in
the destruction of eggs, cysts, and larvae of many para-

sites. Miller, Chi-Rodriguez, and Nichols (1961) studied
several dung beetles which had ingested human dung
containing eggs of hookworm, Ascaris, and whipworm,
and cysts of Entamoeba coli, Endolimax, and Giardia.
No eggs and few cysts were recovered from Canthon pi-
lularius (L.) and C. vigilans Lec., and no Ascaris were
found in C. pilularius after the ingestion of embryon-
ated eggs. Unaltered whipworm eggs and cysts, but no
hookworm and few Ascaris eggs were recovered from
Phanaeus vindex MacL. and P. igneus MacL. Our
largest dung beetle, Dichotomius carolinus (L.), regu-
larly contained and excreted ingested eggs. These
authors concluded that "Although dung beetles may
be significant in other ways in dissemination of hook-
worm, the species most abundantly attracted to human
stools destroy ingested eggs of this parasite and of
Ascaris and do not transport them internally."

(Only subfamilies I-VI are treated in Part I)

Subfamily I. Scarabaeinae
Tribe I. Scarabaeini
Genus 1. Deltochilum Eschscholtz
gibbosum gibbosum (Fabricius)

Genus 2. Canthon Hoffmansegg
chalcites (Haldeman)
pilularius (Linnaeus)
vigilans LeConte

Genus 3. Boreocanthon Halffter
depressipennis (LeConte)
probus (Germar)

Genus 4. Melanocanthon Halffter
bispinatus (Robinson)
granulifer (Schmidt)
punctaticollis (Schaeffer)

Genus 5. Glaphyrocanthon Martinez
viridis viridis (Beauvois)

Genus 6. Pseudocanthon Bates
perplexus (LeConte)

Tribe II. Coprini

Genus 1. Ateuchus Weber
histeroides histeroides Weber
lecontei (Harold)

Genus 2. Dichotomius Hope
carolinus (Linnaeus)

Genus 3. Copris Muller
gopheri Hubbard
howdeni Matthews and Halffter
inemarginatus Blatchley
minutus (Drury)

Genus 4. Phanaeus MacLeay
igneus igneus MacLeay
igneus floridanus d'Olsoufieff
index vindex MacLeay

Tribe III. Onthophagini
Genus 1. Onthophagus Latreille
aciculatulus Blatchley
concinnus Laporte
depressus Harold
hecate blatchleyi Brown
oklahomensis Brown
orpheus orpheus (Panzer)
pennsylvanicus Harold
polyphemi polyphemi Harold
polyphemi sparsisetosus Howden and
striatulus floridanus Blatchley
subaeneus (Beauvois)
tuberculifrons Harold

Tribe IV. Oniticellini
Genus 1. Oniticellus Serville
*cubiensis Laporte

Subfamily II. Aphodiinae
Tribe I. Aphodiini

*Those species preceded by an asterisk are questionably
recorded from Florida; for further details, see the in-
dividual species discussions.

Genus 1. Aphodius Illiger
aegrotus Horn
bicolor Say
campestris Blatchley
crassulus Horn
cuniculus Chevrolat
fimetarius (Linnaeus)
floridanus Robinson
granarius (Linnaeus)
haemorrhoidalis (Linnaeus)
haldemani Horn
laevigatus Haldeman
*lentus Horn
lividus (Olivier)
lutulentus Haldeman
parcus Horn
rubeolus (Beauvois)
*rusicola Melsheimer
stercorosus Melsheimer
stupidus Horn
troglodytes Hubbard

Tribe II. Eupariini

Genus 1. Euparia Serville
castanea Serville

Genus 2. Myrmecaphodius Martinez
excavaticollis (Blanchard)

Genus 3. Ataenius Harold
abditus (Haldeman)
alternatus (Melsheimer)
apicalis Hinton
brevicollis Wollaston
brevinotus Chapin (n.sp., near)
cylindrus Horn
erratus Fall
exiguus Brown
fattigi Cartwright
figurator Harold
gracilis (Melsheimer)
havanensis Balthasar
imbricatus (Melsheimer)
insculptus Horn
integer Harold
languidus Schmidt
miamii Cartwright
ovatulus Horn
picinus Harold
platensis Blanchard
rhyticephalus Chevrolat
rudellus Fall
saramari Cartwright
simulator Harold
spretulus (Haldeman)
strigatus (Say)
waltherhorni Balthasar
wenzelii Horn
n. sp. #1
n. sp. #2
n. sp. #3

Genus 4. Pseudataenius Brown
n. sp. (near socialis Horn)
Genus 5. Aphotaenius Cartwright
carolinus (Van Dyke)
Tribe III. Psammodiini
Genus 1. Psammodius Fallen
armaticeps Fall
bidens Horn
cruentus Harold
malkini Cartwright
Genus 2. Pleurophorus Mulsant
longulus Cartwright
micros (Bates)
Genus 3. Rhyssemus Mulsant
*scaber Haldeman

Subfamily III.
Genus 1.

Subfamily IV.
Genus 1.

Hybosorus MacLeay
illigeri Reiche

Ochodaeus Serville
frontalis LeConte
musculuss Say

Subfamily V. Geotrupinae
Tribe I. Bolboceratini
Genus 1. Bolboceras Kirby
floridensis (Wallis)
Genus 2. Bradycinetulus Cockerell
ferrugineus (Beauvois)
Genus 3. Bolbocerosoma Schaeffer
hamatum Brown
Genus 4. Eucanthus Westwood
alutaceus Cartwright
impressus Howden
subtropicus Howden
Tribe II. Geotrupini
Genus 1. Geotrupes Latreille
blackburnii blackburnii (Fabricius)
egeriei Germar
Genus 2. Peltotrupes Blanchard
profundus Howden
young Howden
Genus 3. Mycotrupes LeConte
cartwrighti Olson and Hubbell
gaigei Olson and Hubbell
pedester Howden

Subfamily VI. Acanthocerinae
Genus 1. Acanthocerus MacLeay
aeneus MacLeay
Genus 2. Cloeotus Germar
aphodioides (Illiger)
globosus (Say)

Subfamily VII. Melolonthinae
Tribe I. Sericini
Genus 1. Serica MacLeay
aemula Dawson
aspera Dawson
atracapilla (Kirby)
delicate Dawson
floridana Dawson
frosti Dawson
georgiana georgiana Leng
georgiana lecontei Dawson
intermixta Blatchley
iricolor (Say)
panda Dawson
parallel Casey
peleca Dawson
pusilla Dawson
rhypha Dawson
sericea (Illiger)
spicula Dawson
tantula Dawson
vespertina (Gyllenhall)
Tribe II. Liparetrini
Genus 1. Hypotrichia LeConte
spissipes LeConte
Tribe III. Melolonthini
Genus 1. Diplotaxis Kirby
bidentata LeConte
frontalis LeConte
languida LeConte
liberta (Germar)
punctatorugosa Blanchard
rufa Linell
subcostata Blanchard
subcostata n. ssp.
Genus 2. Polyphylla Harris
gracilis Horn
occidentalis (Linnaeus)
pubescens Cartwright
Genus 3. Phyllophaga Harris
aemula (Horn)
bruneri Chapin
*calceata (LeConte)
clemens (Horn)
clypeata (Horn)
crenulata (Froelich)
cupuliformis Langston
debilis (LeConte)
diffinis (Blanchard)
dispar (Burmeister)
elizoria Saylor
elongata (Linell)
ephilida (Say)
floridana Robinson
forsteri (Burmeister)
glaberrima (Blanchard)
hirticula (Knoch)
ilicis (Knoch)
infidelis (Horn)
knochii (Schoenhert and Gyllenhal)
latifrons (LeConte)

lota Luginbill
luctuosa (Horn)
marginalis (LeConte)
mariana Fall
micans (Knoch)
okechobea Robinson
ovalis Cartwright
parvidens (LeConte)
profunda (Blanchard)
prununculina (Burmeister)
quercus (Knoch)
schaefferi Saylor
submucida (LeConte)
subpruinosa (Casey)
tecta Cartwright
tristis (Fabricius)
ulkei (Smith)
uniforms (Blanchard)
young Cartwright
new species (Eustis)
new species (Chumukla)
Genus 4. Plectris Serville
aliena Chapin
Tribe IV. Pachydemini
Genus 1. Gronocarus Schaeffer
autumnalis Schaeffer
multispinosus Howden
Tribe V. Macrodactylini
Genus 1. Macrodactylus Latreille
subspinosuss (Fabricius)
Tribe VI. Hopliini
Genus 1. Hoplia Illiger
equina LeConte
floridana Fisher
meridionalis Boyer
Subfamily VIII. Rutelinae
Tribe I. Anomalini
Genus 1. Anomalepta Casey
flaccida Casey
semilivida (LeConte)
Genus 2. Anomala Samouelle
exiguua (Schwarz)
flavipennis Burmeister
innuba (Fabricius)
ludoviciana Schaeffer
minute Burmeister
nigropicta Casey
parvula Burmeister
undulata Melsheimer
Genus 3. Pachystethus Blanchard
floridana Robinson
marginata (Fabricius)
oblivia (Horn)
Genus 4. Strigodermella Casey
pygmaea (Fabricius)
Tribe II. Rutelini
Genus 1. Pelidnota MacLeay
lutea (Olivier)
punctata (Linnaeus)
Genus 2. Cotalpa Burmeister
lanigera (Linnaeus)

Genus 3. Rutela Latreille
formosa Burmeister
Genus 4. Parastasia Westwood
brevipes LeConte
Subfamily IX. Dynastinae
Tribe I. Cyclocephalini
Genus 1. Dyscinetus Harold
morator (Fabricius)
Genus 2. Cyclocephala Latreille
borealis Arrow
immaculate (Olivier)
miamiensis Howden and Endrodi
parallel (Casey)
puberula (LeConte)
Tribe II. Oryctini
Genus 1. Euetheola Bates
rugiceps (LeConte)
Genus 2. Bothynus LeConte
cuniculus (Fabricius)
morio LeConte
neglectus LeConte
subtropicus (Blatchley)
Genus 3. Aphonus LeConte
castaneus (Melsheimer)
variolosus (LeConte)
Genus 4. Strategus Hope
antaeus (Drury)
julianus Burmeister
splendens (Beauvois)
Tribe III. Dynastini
Genus 1. Dynastes Kirby
tityus (Linnaeus)

Tribe IV. Phileurini
Genus 1. Phileurus Latreille
castaneus Haldeman
truncatus (Beauvois)
Subfamily X. Cetoniinae
Tribe I. Gymnetini
Genus 1. Cotinis Burmeister
nitida (Linnaeus)
Tribe II. Cetoniini
Genus 1. Euphoria Burmeister
inda (Linnaeus)
limbalis Fall
sepulchralis (Fabricius)
Genus 2. Stephanucha Burmeister
areata (Fabricius)
*thoracica Casey
Tribe III. Cremastocheilini
Genus 1. Cremastocheilus Knoch
*canaliculatus Kirby
*harrisi Kirby
squamulosus LeConte
Tribe IV. Trichiini
Genus 1. Trigonopeltastes Burmeister
delta (Forster)
floridana (Casey)
Genus 2. Trichiotinus Casey
lunulatus (Fabricius)
piger (Fabricius)
rufobrunneus Casey
Tribe V. Valgini
Genus 1. Valgus Scriba
*canaliculatus (Fabricius)


Key to the Florida subfamilies of Scarababaeidae

1. Abdominal spiracles situated in the membrane
connecting the dorsal and ventral corneous
plates, the last spiracle being covered by the
elytra (Fig. 3); ligula always separate from
the mentum (visible only after dissection);
tarsal claws, when present, simple; antennal
club always of three segments; primarily dung
feeding species.......... LAPAROSTICTI-2
1'. Abdominal spiracles partly situated in the su-
perior portion of the ventral plates, the last
spiracle usually visible behind the elytra (Fig.
2,4); ligula sometimes free, usually connate
with the mentum (visible only after dissection);
tarsal claws always present and often toothed;
antennal club sometimes with more than three
segments; primarily plant feeding species....
................... PLEURdSTICTI-7

2(1). Body globular, capable of contracting into a
ball (Fig. 43); middle and posterior tibiae
flattened and dilated (Fig. 392-395); abdomen
with 5 visible ventral segments (Fig. 39); an-
tennae 10 segmented .... ACANTHOCERINAE
2'. Body not globular nor capable of contracting
into a ball; middle and posterior tibiae never
flattened nor dilated (Fig. 41-42, 50-51); ab-
domen with 6 visible ventral segments (Fig.
40); antennae 8-11 segmented ............3
3(2'). Antennae 11-segmented (Fig. 53-54); man-
dibles prominent from above; posterior tibiae
with 2 apical spurs ........ GEOTRUPINAE
3'. Antennae 8-10 segmented; mandibles prominent
or hidden from above; posterior tibiae with 1 or
2 apical spurs ...........................4

4(3'). Posterior tibiae with a single apical spur (Fig.
50-51, 58) (except for Melanocanthon, in
which case the clypeus is quadridentate); pygi-
dium exposed; mandibles hidden from above
............... ......... SCARABAEINAE

4'. Posterior tibia with two apical spurs (Fig. 41-
42, 55-57); clypeus bidentate or without
teeth, never quadridentate; pygidium exposed
or hidden; mandibles hidden or prominent from
above .. ............................. . 5

5(4'). Mandibles not visible from above (Fig. 178,
212, 220, 222, 290, 292-293, 306, 309); an-
tenna 9-segmented; epimera of metathorax
covered ................... . APHODIINAE

5'. Mandibles prominent, visible from above (Fig.
310, 319, 321-23); antennae 10-segmented;
epimera of metathorax covered or visible.... 6

6(5'). Body densely pubescent dorsally (Fig. 321);
antennal club with 3 normal segments, the first
not excavated for reception of the second; color
light yellow to brown; longer middle tibial spur
pectinate (Fig. 324-26, 330).. OCHODAEINAE

6'. Body glabrous, shining dorsally (Fig. 310); an-
tennal club with the first segment hollowed
for reception of the second which is nearly con-
cealed (Fig. 317-18); color black; both mid-
dle tibial spurs entire.......HYBOSORINAE
................. . Hybosorus illigeri Reiche

7(1'). Tarsal claws unequal; posterior tibiae with 2
apical spurs; labrum visible from above......
. . . . . . . . . . . . .... RUTELINAE
7'. Tarsal claws usually equal (if unequal, then
posterior tibiae without apical spurs); labrum
visible or hidden from above .............8
8(7'). Mandibles concealed by the clypeus; antennae
7-10 segmented; tarsal claws usually cleft, bifid,
or toothed (Fig. 48) not more than one pair
of abdominal spiracles exposed below edges of
elytra; color yellow-brown to black, never metal-
lic or green ............ MELOLONTHINAE
8'. Mandibles usually visible from above; antennae
9-10 segmented; tarsal claws variable, often not
cleft or toothed; usually at least two pair of
abdominal spiracles exposed below edges of
elytra; color variable, often with metallic green
. . . . . . . . . . . . . . . . . . . . . . . . . . . .. .. . . . . . . . . 9
9(8)'). Front coxae transverse, not prominent; mandi-
bles bent, expanded and leaf-like, often
notched; head and/or pronotum armed with
horns or protuberances; scutellum as wide as
long, rounded behind; color brown to black,
never metallic nor green ...... DYNASTINAE
9'. Front coxae conical, prominent; mandibles not
bent or leaf-like; head and/or pronotum rarely
with horns or protuberances; scutellum longer
than wide, pointed behind; color variable but
often metalilc and/or green . . .CETONIINAE

(Fig. 1, 81, 84, 100, 107, 122, 125, 128,
133, 136, 145, 155, 177)

TYPE GENUS: Scarabaeus Linnaeus, 1758:
345 (by autonomyy.

DIAGNOSIS: Variable in form and size (length 2 to
30mm), but usually oval and rounded. Color variable
from shining metallic green, blue or bronze, to dull,
matte black or brown; rarely with spots or maculations
of red to orange (in two species of Onthophagus).
Mouthparts primarily membranous, only the outer
margin of the mandibles corneous. Clypeus expanded,
concealing the mouthparts from above. Antennae eight
or nine segmented; club always three segmented. Epi-
mera of metathorax covered; mesosternum very short;
middle coxae oblique, widely separated; posterior tibia
with a single spur (except in Melanocanthon which
has two). Pygidium exposed and usually triangular in
shape. Six visible abdominal segments.
In some genera (Phanaeus and Deltochilum) the
anterior tarsi are usually missing, a feature not found
in other subfamilies. In some species the males have
well developed horns on the head and protuberances

and excavations on the pronotum (never in the tribe
TAXONOMIC NOTES: This is a large subfamily of
several thousand species (no accurate count available)
representing 5 tribes, 14 subtribes and 201 genera
(Halffter and Matthews, 1966). The group as a whole
is fairly well known, with modern revisions in several
genera. The status of the classification has been
summarized by Halffter and Matthews as follows:
While in some groups-those which have
been revised by recent workers with modern
concepts-knowledge is almost at the beta level,
in certain others the generic concepts are very
confused and no real progress has been made
since the time of Harold, 100 years ago. In the
first (beta) category we have the Oniticellini,
Onitini, Gymnopleurina, some of the Scarabaeina,
and the Madagascan and American Canthonina.
The latter-the group of the subfamily in which
the most work is being done at present-are in
the process of revision, and their classification

is approximating a phylogenetic scheme. All of
the American canthonine genera included in our
list represent natural groups of species, al-
though in some cases it is likely that their rank
will be lowered to the level of subgenera in the
future. At the other extreme, among groups at
the lowest alpha level of study, we have the
remaining Canthonina, some of the Coprina, and
especially the Dichotomina.
All of the tribes (except Onitini: Oriental, Ethio-
pian, and Palearctic) are represented in Florida, al-
though the single record of the Oniticellini is doubtful.
Only 5 of the 14 subtribes are represented (i.e., Oniti-
cellina, Dichotomina, Phanaeina, Coprina, and Can.
thonina) by 12 genera with about 36 species and
The name Coprinae has been used for this group
by many authors (e.g., Gillet, Harold, and Peringuey).
It has been called Coprophagi (Latreille, 1802; Laporte,
1840); Coprophaga (Burmeister, 1842), and Coprides
(Erichson, 1847; Lacordaire, 1856). It was considered
a full family by Balthasar (1963) and by some other
European authors. However, recent American authors
have usually considered it of subfamily rank, and they
have used the name Scarabaeinae (e.g., Arnett, 1962;
Halffter and Matthews, 1966).
BIOLOGY: Nearly all members are dung feeders in
both the adult and larval stages, but a few are necro-
phagous or saprophagous. Although practically none
of the species is harmful, the group has been the sub-
ject of considerable interest for centuries; beginning
with their worship by the ancient Egyptians, through
the early classic behavior studies by the French nat-
uralist Fabre, to the wave of interest of the present
The literature on the biology of dung beetles is
therefore quite extensive, much of it appearing in
scattered journals and in numerous languages. This
situation has long been a handicap to anyone working
on the group, but it has been alleviated by the recent
monumental review, in English, by Halffter and Mat-
thews (1966). They have reviewed the entire literature
on the biology of Scarabaeinae, along with consider-
able original data on the subject. This publication is
the most significant modern contribution to the biology
within the family Scarabaeidae, and should greatly
enhance future work and create considerable interest
in dung beetles. Therefore, I will not dwell on the
subject further here, but the reader is urged to read
the above paper for a thorough and enlightening ac-
count of the biology of this fascinating subfamily.

Key to the Florida tribes and genera of
Scarabaeinae (=Coprinae)
1. Middle and posterior tibiae slender, curved,
scarcely enlarged at the apex (Fig. 51);
head and pronotum without horns or pro-
tuberances; elytral striae obsolete, poorly de-
fined .................. (Scarabaeini)--4

1'. Middle and posterior tibiae enlarged at the
apex (Fig. 50); head and pronotum often
with horns and/or protuberances; elytral
striae obvious or obsolete ..............2
2(1'). Third segment of labial palpi distinct; ely-
tral striae distinct, or if not, the intervals
reticulately punctate; color black or bronze,
elytra never maculate; length 5-29 mm....
.................... . (Coprini)- 9
2'. Third segment of labial palpi inconspicuous;
elytral striae obsolete; color black to metallic
green, the elytra sometimes maculate with
small orange to red spots; length 2-8 mm..
. . . .. . . .. . .. . . . . .. . .. . . . . . .. . . . . . . . . 3
3(2'). Antennae 9-segmented; scutellum hidden; sev-
eral common Florida species; length 2-8 mm;
(Fig. 155) . .. (Onthophagini)-Onthophagus
3'. Antennae 8-segmented; scutellum small but
visible; single Florida record doubtful; length
5-7 mm; (Fig. 177) ......... (Oniticellini)
................ Oniticellus cubiensis Lap.


Anterior tarsi absent (Fig. 52); elytral epi-
pleural fold broad; male with enlarged
"hump" on anterior one-third of elytra;
length 20-25 mm; (Fig. 81) ..............
.......... Deltochilum g. gibbosum (Fab.)

4'. Anterior tarsi present; elytral epipleural fold
very narrow or absent; neither sex with en-
larged hump on elytra; length 2-22 mm...5


Posterior tibia with two terminal spurs; (Fig.
57, 107) ................. Melanocanthon

5'. Posterior tibia with a single terminal spur
(Fig. 58) ............................ 6


First segment of posterior tarsus equal to or
longer than the second; surface dull, granu-
lar, or shining; color gray to bronze, if
green then dull granulate, not shining; cly-
peus bidentate or quadridentate; length 5-22
mm ............................ ... 7

6'. First segment of posterior tarsus shorter than
the second; surface shining, never granulate
(although minutely pebbled); color usually
bright green, shining, but sometimes purple
bronze, never gray or black; clypeus biden-
tate; length 2-4 mm; (Fig. 122) .........
.......... Glaphyrocanthon viridis (Beauv.)
7(6). Base of pygidium without margin; clypeus
quadridentate; dorsal surface shining not
granulate; color bronze to black with green
sheen; length 3-4 mm; (Fig. 125) .......
........... Pseudocanthon perplexus (Lec.)
7'. Base of pygidium margined, clypeus bidentate
or quadridentate; dorsal surface granulate,
not shining; color dull gray to green; length
4-22 mm; (Fig. 84, 100) ............. 8

8(7'). Clypeus bidentate; anterior border of pos-
terior femur margined; length 11-22 mm;
(Fig. 84, 105) .................. Canthon

8'. Clypeus quadridentate; anterior border of
posterior femur without margin; length 4-10
mm; (Fig. 100, 106) ........ Boreocanthon

9(2). Anterior coxae very transverse, not promi-
nent; head and pronotum without horns or
protuberances; color bronze to black, never
bright green; length 5-7 mm; (Fig. 128) ....
............ .................. A teuchus

9'. Anterior coxae short, conical and prominent;
head and pronotum often with horns or pro-
tuberances; color black to bright metallic
green; length 8-29 mm ............... 10

10(9'). Anterior tarsi absent in males (Fig. 49),
the females lacking claws on these tarsi; color
metallic green or blue, often with reddish
reflections; elytral striae not prominent, the
intervals reticulately punctate; (Fig. 1, 145).
....................... Phanaeus
10'. Anterior tarsi and claws present in both
sexes; color black, never green or metallic;
elytral striae deeply impressed, the intervals
smooth, convex ...................... 11
11(10'). Elytral striae seven; clypeus not notched
medially; length 20-30 mm; (Fig. 133).....
.............. Dichotomius carolinus (L.)
11'. Elytral striae eight; clypeus usually notched
medially; length 8-15 mm; (Fig. 136) .....
................................. Copris

(Fig. 57-58, 81, 84, 100, 105-107, 122, 125)

This tribe was divided into seven subtribes by
Halffter and Matthews (1966), but only one (Cantho-
nina) is represented in Florida. This subtribe was
treated as a tribe, the Canthonini, by Vulcano and
Pereira (1964). It was called Canthonides by Paulian
(1938), Gillet (1911), Peringuey (1901), and Black-
welder (1944). LeConte and Horn (1883) treated our
species under the tribe Coprini and subtribe Ateuchini.
The American species have received considerable
attention recently (e.g., Martinez, Pereira, Halffter,
Matthews). The subtribe Canthonina was recently cata-
logued for the western hemisphere (Vulcano and
Pereira, 1964). In this catalogue 39 genera are listed,
of which the following 6 occur in Florida: Boreocan-
thon, Melanocanthon, Canthon, Deltochilum, Glaphyro-
canthon, and Pseudocanthon.
All of the Florida species, except Deltochilum gib-
bosum Fab., have been listed in the genus Canthon
in previous papers on U. S. species. Arnett (1962), in
his "Beetles of the United States," does not recognize
the genera Boreocanthon, Glaphyrocanthon, Melanocan-
thon, and Pseudocanthon. The process of fragmenting
the old genus Canthon is still underway, and generic
concepts are not firmly established. Two of the U. S.
workers (Howden and Matthews) have raised some
doubts about the validity of several recently described
genera (e.g., Boreocanthon and Nesocanthon). There
undoubtedly will be considerable shuffling of names
for a few years. The entire subtribe appears to have
its center of orign in the American tropics, with only
a few species entering the U. S. Howden (1966b), ex-
pressed doubt about the validity of certain genera, but
he admitted that he ". . . cannot properly assess .. ."
them at this time. Therefore, I am inclined to follow
the treatment derived by the workers in South America
and Mexico.
The tribe is characterized by the slender, often
curved, middle and posterior tibiae. 'he dorsal surface

is often granular, and the elytral striae are poorly
defined. The head and pronotum are. both without
horns or protuberances, although the clypeus is biden-
tate or quadridentate. The posterior legs are often
elongate and used in rolling balls of dung. The size
range is from 2 mm (Glaphyrocanthon) to 22 mm.
All of our species are coprophagous as adults and
larvae. The adults roll and bury balls in which a single
egg is laid, thus providing food for the larvae. Details
of the nidification behavior probably vary somewhat in
our species, but in general it may be similar to that
reported for Canthon pilularius (L.) (Matthews, 1963).
These beetles are familiar to most farm boys and are
called "tumble bugs."

Genus DELTOCHILUM Eschscholtz
(Fig. 81)

Deltochilum Eschscholtz 1822:37.
Anamnesis Vigors 1826:510.
Hyboma Serville 1828:352 (not Hyboma Huebner,
Deltachilum Esch., LeConte 1863b:36 (misspelling).
Meghyboma Kolbe 1893:192.
Annamesis Vigors, Gemminger and Harold 1869:995
(misspelling of Anamnesis).
TYPE SPECIES: D. (Deltochilum) dentipes Esch-
scholtz 1822:38 (by monotypy).
DIAGNOSIS: Large for the tribe (length to 25 mm,
width to 19 mm); dull black; anterior tarsi lacking;
middle and posterior tibiae long, slender, the posterior
ones bent inward abruptly at the middle, scarcely en-
larged at the tip. Sexual dimorphism pronounced:
males with prominent swellings tumescencess) on the

anterior one-third of the elytra; posterior tibiae more
strongly bent; anterior tibiae with a median tooth
projecting ventrally, more strongly developed in the
male. Clypeus quadridentate; inner pair of teeth acute
with a carina extending from the tip back onto the
clypeus; outer pair obtusely angulate, not prominent.
Inner pair of teeth separated by nearly 2 mm, arcuately
emarginate between. Elytra descending abruptly near
apex with three to five variably shaped (mostly longi-
tudinal) carinae at this point. Humeral carina short
and longitudinal. Epipleural fold broad (unlike other
Canthonini), the elytra bordered laterally by a sharp
carina. Surface finely alutaceous, matte, shining only
at tumescences, posterior elytral carinae, and humeral
carinae. Normally dull black, occasionally with a faint
dark purple caste; never metallic. Museum specimens
often "greased" and encrusted with soil or carrion.
Middle tibia with two apical spurs; posterior tibia
with one apical spur. Anterior tibia tridentate, anterior
two teeth nearer each other than the posterior one.
Metasternum anteriorly depressed at middle in male
only. Pygidium large, flattened, not noticeably convex,
nearly vertically oriented.
TAXONOMIC NOTES: In a recent catalogue of the
Canthonini, Vulcano and Pereira (1964) listed 73
species in the genus which is divided into 9 subgenera
as follows: Aganhyboma (4), Calhyboma (11), Euhy-
boma (1), Telhyboma (1), Hybomidium (7), Parahy-
boma (2), Rubrohyboma (1), Deltohyboma (43), and
Deltochilum (3). Howden (1966b) added two new sub-
species, one new species, and synonymized one species
to bring the total to 75 species and subspecies cur-
rently recognized. The synonymy cited above is mostly
from Vulcano and Pereira (1964).
Our single species, D. gibbbosum (Fab.), is the
ype of the subgenus Hybomidium Shipp (1897:195).
his name was proposed to replace Hyboma Serville
(1828:352) which was preoccupied by Hyboma Hueb-
ner (1820:200) in the Lepidoptera. Paulian (1938:
259) subsequently described the new subgenus Tetrao-
dontides with D. gibbosum as the type species, al-
though in the same paper (p. 238) he listed D.
gibbosum as the type species of Deltochilum. Since D.
dentipes Esch. was previously selected as type of the
genus, and Tetraodontides is a synonym of Hybomid-
ium, our species becomes Deltochilum (Hybomidium)
gibbosum (Fab.)
almost exclusively Neotropical, with only two species
entering the United States. Of these two, D. scabrius-
culum Bates is known from Bronwsville, Texas, south
to Guatemala and Costa Rica. The other, D. gibbosum
(Fab.), is known from Kentucky south to Florida and
west to Texas, with disjunct subspecies from Panama
and Mexico. Ten of the species and subspecies occur
in Mexico and Central America, with the remainder
of the 75 being South American. The origin of the
genus appears to be northern South America, possibly
coincidental with that of the genus Canthon, for which
Halffter (1961) postulated "Arquibrazil."
BIOLOGY: This genus contains the largest species of
North American "tumble bugs." As the name implies,

they roll balls of food material which are used pri-
marily for egg deposition and in which the developing
larvae feed. It is not known if some of these balls are
also used as adult food as they are with some species
of Canthon. The food materials are quite varied and
include nearly any kind of decaying plant or animal
substance. However, there appears to be a decided
preference for carrion. Specific foods observed are:
dead crabs, chicken feathers, dog carrion, decaying
cantaloupe, decaying fish, human dung, horse dung,
fermenting malt, and decomposing fungi. Walker
(1957) found D. gibbosum at decaying fish and decay-
ing cantaloupe in the three forested habitats investi-
gated, but none was taken on the same baits employed
in an old field. Many of the species seem to be more
abundant in wooded areas, but at least some species
are common in semi-arid regions (Howden, 1966b).
Fungi do not appear to be attractive except in the
late stages of decomposition. Blatchley (1928a:62)
mentioned several specimens of D. gibbosum in a "...
putrid, extremely foetid mass of fungi.", and Howden
and Ritcher (1952) stated that fungi seem attractive
only in the fall. Walker (1957) reported that D. gib-
bosum was attracted to dead fish within one day
after the fresh bait was deposited, but cantaloupe was
not attractive until after the 7th day when the flesh
was "... very soft, much liquid." My own experiences
in Mexico and Central America indicate that, although
the habitats are variable, specimens are rarely en-
countered in open pastures. Gibson (In Howden, 1966b:
738) found D. scabriusculum montanum Howden active
around horse dung, but none was utilizing readily
available cow dung nearby. At least some of the
species are attracted to lights.
The brood balls, in which eggs are deposited, are
not round as are those of most species of Canthon.
They have been described as "narcissus-bulb" or "pear-
shaped," although they have been discovered for very
few species. Their shape implies that they are rolled
in a spinning fashion rather than over and over like a
ball as is the case in Canthon. The posterior tibiae are
especially suited for this task. It is not known if both
sexes participate in construction and/or rolling and
deposition. The sexual dimorphism of the posterior
tibiae (males more curved, Fig. 51) is a possible
reflection that the male plays a more active role than
the female. The balls are often made of the food
material with dirt and leaves packed on the outside.
Their general appearance is quite unlike that of the
smooth, round ball of Canthon. They are most fre-
quently deposited at shallow depths near some natural
feature such as a rock or log. Howden and Ritcher
(1952) reported a two-inch deep, circular, cup-shaped
depression made by the adult, in which a brood ball
was found. These balls are the largest known to be
constructed by New World dung beetles (this is exclud-
ing the dung plugs formed by species of Dichotomius
and Phanaeus) and they often exceed two inches in
diameter and height. The balls of our related Canthon
rarely exceed three-fourths of an inch in diameter.
The behavior has not been observed in any detail but
should provide some interesting comparative data
when it is recorded.

The larvae of few species are known, but they are
probably all similar in general appearance. Those
known are large, gray-white, "hump-backed" grubs
similar to other genera of Scarabaeinae. The most
useful taxonomic character for separating them from s
other genera appears to be the setal pattern of the
venter of the last abdominal segment. Larval develop-
ment is very rapid in those species which have been
studied, taking as little as 20 days from egg to third )
instar (Howden and Ritcher, 1952).
SELECTED REFERENCES: Blatchley, 1928a:62;
Cartwright, 1949b:38; Howden and Ritcher, 1952:53-
57; Howden, 1966b:733-740; Halffter and Matthews,
Deltochilum gibbosum Fob. - ---
1966 (numerous pages referring to biological notes on
15 species); Paulian, 1938:296; Pereira and Martinez,
1956:120-125; and Vulcano and Pereira, 1964:639-660.

Deltochilum gibbosum gibbosum
(Fig. 81-83)
Scarabaeus gibbosus Fabricius 1775:28.
Copris gibbosus (Fab.), Olivier 1790:141, 172.
Ateuchus gibbosus (Fab.), Fabricius 1801:57. 8
Hyboma gibbosa (Fab.), Serville 1828:353.
Deltochilum gibbosum (Fab.), Burmeister 1848:134.
Deltachilum gibbosum (Fab.), LeConte 1863b:36 (mis-
Deltochilum g. gibbosum (Fab.), Bates 1887:36.
DIAGNOSIS: Easily distinguished from other Florida
Canthonini by the lack of anterior tarsi, broad epi-
pleural fold, large size (up to 25 mm in length),
shape and position of the clypeal teeth, anterior elytral
tumescences of the male, carinate elytral margins, and
the longitudinal humeral carina.
TAXONOMIC NOTES: Typical gibbosum is apparently
confined to the southeastern U. S. Howden (1966b:
736) stated, ". .. the species is represented by a com-
plex of related forms extending through Mexico and
Central America into South America. Several seemingly
disjunct populations show constant differences, but, be-
cause of the paucity in collections of specimens from
critical areas, these differences are difficult to assess.
For the present I have treated the various populations
as subspecies, partly to indicate their close relation-
ship, and partly to indicate the possibility of inter-
* breeding." He recognized two additional subspecies:
sublaeve Bates from Mexico and panamensis Howden
from Panama.
Recorded from Alabama, Georgia, Louisiana, Missis-
sippi, North Carolina, South Carolina, Tennessee,
Texas, and I have also seen specimens from southeast-
ern Kentucky (new state record). In Florida it occurs
from Pensacola to Big Pine Key, the spotty records
probably reflecting its secretive habits, and it prob-
ably occurs throughout the state.
BIOLOGY: (see remarks under the genus). This spe-
Fig. 81, Deltochilum g. gibbosum (Fab.), line = 8 mm. cies is probably much more abundant than the records


would indicate, but it is secretive in its habits. It is
possible that it is primarily active at night (as is
Canthon vigilans). Specimens are readily attracted to
chicken feathers, especially in forested areas. I have
taken two dead specimens apparently trapped in a box
in which dead blue crabs had been deposited. Other
hosts include dog carrion, fermenting malt, Japanese
beetle trap, decaying fish, decaying cantaloupe, human
dung, and decomposing fungi. There is a single record
at light (Frost, 1964:142). In Florida it apparently
occurs throughout the year, with records from January
through November. No behavioral observations have
been recorded.

SPECIMENS EXAMINED: 85, of which 70 were
from 22 Florida localities (for complete data see Ap-
pendix 1).

SELECTED REFERENCES: Angell, 1913:169, Blatch-
ley, 1920b:43, 1828a:62; Cartwright, 1949b:38; Frost,
1964:142; Hebard, 1903:261; Howden, 1966b:736, Fig.
17-18; Howden and Ritcher, 1952:53, 11 Fig.; Paulian,
1938:259-262, Map; 1939:Fig. 1, 10; Pereira and Mar-
tinez, 1956:125; Vulcano and Pereira, 1964:648-649;
Walker, 1957:Table 5, 7, Fig. 6, 9-11.

Genus CANTHON Hoffmannsegg
(Fig. 84-99)

Canthon Hoffmannsegg 1817:38.
Coprobius Latreille 1829:535.
Coeloscelis Reiche 1841:213.

TYPE SPECIES: Scarabaeus pilularius Linnaeus
1758, by subsequent designation of Paulian (1938-

DIAGNOSIS: Typical Scarabaeinae; medium sized (10-
22 mm long); at least part of dorsal surface granular;
clypeus bidentate; posterior femur margined anteriorly;
pygidium margined at base; anterior tarsi present;
middle and posterior tibiae slender, scarcely enlarged
at tip; sexual dimorphism not noticeable; head and
pronotum unarmed, without horns or protuberances:
color black to brown, greenish to bronze but never
shiny metallic. Anterior part of the prothorax below
not excavated to receive the fore femora, and no
transverse carina delimiting this area. Anterior tibial
spur variable, often bifurcate in the male, simply acute
in the female. Middle tibia with two spurs; posterior
tibia with one spur.

TAXONOMIC NOTES: The genus, in its narrow sense,
has been recently monographed for North America
(Halffter, 1961) and catalogued for the world (Vul-
cano and Pereira, 1964). In these papers 98 species
are recognized for the world. Until recent years this
genus was more broadly applied and included all of
the species treated here under the genera GbKphyrocan-
thon, Pseudocanthon, Boreocanthon, and Melanocan.

thon. Numerous other genera have also been split from
the old genus Canthon, especially for the South Amer-
ican species, and this process is still going on. Some
doubts have been raised concerning the validity of
some of these divisions (Matthews, 1966; Howden,
1966b), but they are provisionally accepted here until
a complete revision of the tribe is made (currently
in progress by Halffter and Martinez, 1966-70).

Fig. 84. Canthon pilularius (L.), line = 7mm.

Our Florida species are similar in appearance and
often difficult to determine without comparative ma-
terial. However, the characters mentioned in the key
and the illustrations should serve to distinguish the
species. Characters of the male genitalia are useful but
not greatly different among our species. The internal
sac has been used for distinguishing many of the
South American forms. No subspecies are recognized
for the U. S. species, although several have been de-
scribed for Mexican and Central American species
(Halffter, 1961). Our species vary primarily in color
(from black to blue or green) and size (10 to 22
mm in length). Variation in the labrum of one species
(vigilans) is described as clinal by Halffter (1961).
The Florida species are generally larger than many of
the Central and South American species, and one of
our species (vigilans) attains maximum size for the
genus (22 mm).
the 98 species listed by Vulcano and Pereira (1964)
are Neotropical, with only 8 species being found in
North America. The genus appears to have had its
origin in South America. Four species are recorded
from the Antilles (Matthews, 1966), although the three
previously described species were placed in their new

genus Nesocanthon by Pereira and Martinez, and the
fourth was described by Matthews (1966) as occupying
a ". . . taxonomically highly isolated . . ." position in
the genus. Only three of the eight North American
species occur in the Eastern U. S., and all three have
been recorded from Florida. In Florida, only one
species (pilularius) appears to occupy the entire state,
although I have not personally seen specimens from
south of Miami or from the Keys. Blatchley (1928:61)
listed a single specimen from Key West. Of the re-
maining two species, chalcites is known only from four
isolated localities (Welaka, Ocala Nat. For., Torreya
St. Pk., and Miami), and vigilans probably occupies
the northern two-thirds of the state as far south as
Ft. Myers.

BIOLOGY: These are the so called "tumble bugs"
which are a familiar sight to every farm boy. All of
our species are primarily coprophagous and feed com-
monly in cow and horse dung; although there are some
records indicating a slight trend for sarcophagy. All
of our species roll balls of dung away from the source,
and these may be utilized for adult food (food balls)
or for egg deposition and subsequent larval food
(brood balls). They occur more abundantly in open
pastures and are normally associated with the grass-
land biome. Only one of our species (pilularius) has
been studied in any detail (Matthews, 1963). Only
general behavior, probably common to all of our spe-
cies, will be discussed here. For further detail see the
discussions under each species.
Food balls are constructed, rolled from the source,
and buried by a single beetle of either sex and later
consumed underground. Brood balls are usually more
carefully constructed, sometimes by both sexes, but the
subsequent rolling and burial are executed by the male
only (in pilularius, the only species fully observed);
the female accompanies the male, either walking be-
hind or riding on the ball, but apparently not assisting
in any way. A single egg is laid on each ball and
covered with dung, rendering the ball more pear-
shaped. The brood balls are usually rolled to greater
distances and buried deeper than the food balls. Com-
stock (1940:516) offered a teleological explanation
thusly, ". . . as many predacious insects frequent the
masses of dung from which the balls are obtained, in
order to prey upon the larvae which live there, the
more intelligent tumble bugs remove the food for their
larvae to a safe distance." Little data is available on
the parasites of dung beetles, but such behavior would
seem to offer an evolutionary advantage to the "tumble
bugs." Other stages of this behavior are found within
the genus, from those which have lost (?) the ability
to construct balls, to those which construct balls but
do not roll them from the source (Matthews, 1966).
Certain Central and South American species are
strictly necrophagous.
Most of the species are diurnal, but a few are
strictly nocturnal. One of our species (vigilans) is noc-
turnal, possessing enlarged eyes, and* it is the only
U. S. species attracted to light.

SELECTED REFERENCES: Balthasar, 1939; Blanch-
ard, 1885; Brown, 1928a; Halffter, 1961; Harold,
1868c; Paulian, 1939; Pereira and Martinez, 1956;
Robinson, 1948b; Schmidt, 1922b; Vulcano and Pereira,

Key to the Florida species of Canthon
1. Eyes large for the genus (Fig. 85); body larger
(length 15-22 mm); color uniform black; often
attracted to light............... .vigilans Lee.
1'. Eyes normal for the genus (Fig. 86); body
smaller (length 11-18 mm); color usually
bronze, bluish or greenish, rarely black; never
attracted to light......................... 2
2(1'). Head, pronotum, and pygidium with uniform
minute granules, never with enlarged granules
as are present on the elytra (Fig. 92); uni-
formly bronze (rarely black), never bluish or
greenish; rare in Florida ... .chalcites (Hald.)
2'. Head, pronotum, and pygidium with enlarged
granules similar to those of the elytra (Fig.
93); color variable from bronze to bluish,
greenish, or dull matte gray; common Florida
species ................... . ..pilularius (L.)

Canthon chalcites (Haldeman)
(Fig. 89-92)
Coprobius chalcites Dejean 1836:151 (nomen nudum).
Coprobius chalcites Haldeman 1843:304.
Canthon chalcites Hald., LeConte 1859a:10.
Canthon chalcides Hald., LeConte 1863b:36 (mis-

DIAGNOSIS: Easily distinguished from the two other
Florida species by the lack of coarse granules (like
those of the elytra) on the head, pronotum, and pygi-
dium (Fig. 92). It differs further from vigilans Lee. in
the narrower eyes (as in pilularius). It differs further
from pilularius by its generally larger size (length 13-
21 mm) and uniformly bronze (or rarely black) color.
Anterior tibial spur sexually dimorphic (Fig. 89-91).

TAXONOMIC NOTES: Harold (1868c) synonymized
this species under laevis (a synonym of pilularius),
but he was clearly in error in doing so, as was pointed
out by Horn (1870a). Although the two species are
closely related, the differences in pygidial granulation
are very apparent on comparison of the two. Occa-
sionally a specimen will show only traces of the
normal bronze color, and variability exists also in the
size and shape of the elytral granules. Robinson
(1948b:95) mentioned a form from southern Florida
which had these granules ovate rather than round as in
typical specimens from northern localities. He also
mentioned two specimens from the mountains of
Pennsylvania which had the granules reduced to shin-
ing spots without any height. He further postulated
that these two forms ". . . may prove to be sub-

specific races when more material becomes available."
Halffter (1961) could find little correlation between
the slight amount of variation and geography and thus
did not recognize any subspecies. Although large series
of specimens are available from Missouri and Ne-
braska, this species is rarely collected elsewhere. Until
this material is available in numbers, a proper evalua-
tion of the variability cannot be made.



Tennessee, but it also occurs at sea level in Florida
and in the plains of Kansas, Missouri, and Nebraska.
It has been collected from March through November,
but most records are for June or July. The immature
stages are unknown.
SPECIMENS EXAMINED: 110, of which only 6 were
from Florida as follows: (1) Dade Co., Miami, 12-IX-
34, F. N. Young (USNM); (1) Liberty Co., Torreya
St. Pk., 13-VI-66, H. V. Weems, III; (1) loc. cit., 14-
VIII-68, G. W. Rawson; (1) Putnam Co., Welaka, 1-XI-
39, J. J. Friauf, PH-10 (USNM); (1) Volusia Co.,
Enterprise, III-23 (OSU); (1) Youkon, Fla., Ace.
23983 (USNM). The only other Florida record is that
of Halffter (1961:300) as follows: (3) Marion Co.,
Ocala Nat. For., 21-IX-30, T. H. Hubbell (UMMZ).

89 90 91
Fig. 85-91. Canthon spp.: 85) Dorsal view of the head of
Canthon vigilans Lec. Note the enlarged eyes. 86) Dorsal view
of the head of Canthon pilularius (L.). Note the narrow eyes.
87) Lateral view of the male genitalia of Canthon vigilans
Lec. 88) Lateral view of the male genitalia of Canthon
pilularius (L.). 89) Anterior right tibial spur of male Canthon
chalcites (Hald.) from Miami, Florida. 90) Anterior right
tibial spur of male Canthon chalcites (Hald.) from Oconee
County, South Carolina. 91) Anterior right tibial spur of
female Canthon chalcites (Hald.) from Welaka, Florida.

The type is from "Missouri." Halffter (1961:300) re-
corded it from Alabama, Florida, Georgia, Illinois,
Kansas, Kentucky, Louisiana, Massachusetts, Michigan,
Missouri, Nebraska, New Jersey, North Carolina, Ohio,
Oklahoma, Pennsylvania, South Carolina, Tennessee,
Texas, and Virginia. In Florida it has a very spotty
distribution, although this is probably a reflection of
its rarity. The northernmost record is Torreya State
Park, and the southernmost is Miami. It was not re-
corded in the "Scarabaeidae of Florida" by Blatchley
BIOLOGY: Practically nothing has been published on
the habits of this species. Presumably it is a dung
feeder and rolls balls similar to those of C. pilularius.
I have taken a single specimen in an arbor vitae bog
in Ohio on the carcass of a fox in deep woods. I have
seen a fair series from the mountains of Georgia and

Fig. 92. Stereoscan photo of central area of pygidium of
Canthon chalcites (Hald.) (197X). Note granules similar in size.
- " ''3m _ .

Fig. 93. Stereoscan photo of central area of pygidium of
Canthon pilularius (L.) (197X). Note two sizes of granules.

SELECTED REFERENCES: Blanchard, 1885:166; Dil-
lon and Dillon, 1961:508, P1. 49, Fig. 7; Halffter,
1961:297-301, Fig. 8-9, 71-73; Robinson, 1948b:95, Fig.
3-4; Vulcano and Pereira, 1964:606.

Canthon pilularius (Linnaeus)
(Fig. 84, 86, 88, 93)
Scarabaeus pilularius Catesby 1731-1743: P1. 11 (Pre-
Linnean name).
Scarabaeus pilularius Linnaeus 1758:349.
Scarabaeus laevis Drury 1770:79, P1. 35, Fig. 7.
Scarabaeus hudsonias Forster 1771:24.
Scarabaeus volvens Fabricius 1792:66.
Coprobius obtusidens Ziegler 1844:45.
Canthon laevis viridescens Horn 1870a:47.
DIAGNOSIS: Typical for the genus but generally
smaller (length 12-17 mm) than the other species. Of
the Florida species it is most similar to chalcites, but
differs in the coarse granules of head, pronotum, and
pygidium (Fig. 93). The color is coppery to black in
chalcites, where it is bronze to green or dark blue in
pilularius. It is similar also to vigilans but differs in
the narrower eyes (Fig. 86).
TAXONOMIC NOTES: In the above synonymy I have
listed only the species synonyms in their original
generic combinations. For a fairly complete list of
over 100 citations see Vulcano and Pereira (1964:623-
625). Considerable differences of opinion exist about
the proper name to apply to this species. For many
years it was called laevis Drury in nearly all American

literature. Much of the confusion revolves around the
ambiguous original description of Linnaeus, because
he cited previous references by Catesby (obviously
North America), and he listed the locality as "Amer-
ica." However, he also cited references to pilularius
by Pliny and Aristotle (presumably European and thus
a member of Gymnopleurus and not Canthon). Numer-
ous papers have been written on the subject, and
these are summarized by Halffter (1961).
This species and C. imitator Brown are very
closely related, although distinct species. C. imitator
floridanus Brown was described from Gainesville, Flor-
ida, and later synonymized by Halffter (1961). I am
personally convinced that the subspecies was based on
mislabeled specimens, and imitator, in any of its varia-
tions, does not occur in Florida. The types of florida-
nus and the 180 specimens examined by Halffter
(1961:311) are all labeled: Gainesville, Fla., 1920,
F. W. Walker. I have searched for nearly ten years
for anything resembling imitator around Gainesville,
and, although I have taken several hundred pilularius,
I have found no imitator. I have examined the type
of imitator floridanus in the U. S. National Museum
and part of the series mentioned by Halffter in the
University of Michigan Museum of Zoology and concur
that they are imitator.
The collector of this long series, F. W. Walker,
was normally very meticulous about keeping field notes,
for which a number was usually attached to the la-
beled specimen. Since these specimens do not carry a
specific date nor a field number, their status is imme-
diately questionable. A check of Walker's field notes,
by Dr. T. H. Hubbell and later by myself, revealed
no reference to these specimens. Hubbell stated (in
litt.) that, "I think your surmize that they were mis-

p 5 *

labelled is probably correct, but how it could have
happened is hard to understand. If you are right, the
chances are that Walker had nothing to do with this
The subspecies viridescens Horn was synonymized
by Halffter (1961) and others. The green form on
which it was based is common only in north central
Florida and southern Georgia. The color is quite
variable in specimens taken at a single locality in the
southeast, although those from the north (e.g., Ohio)
are exceptionally uniformly bronze, with no green or
blue reflections. Those from south Florida (e.g., Zolfo
Springs) are uniformly dark blue. Additional studies
should be conducted with this common, wide-ranging
species, in order to ellucidate the status of these color

It apparently occupies nearly the entire area east of
the Rocky Mountains. Previous records from Mexico
are probably all referrable to the closely related imi-
tator. The record of Blanchard (1885:166) for "S. Cal."
is probably also incorrect since no subsequent speci-
mens have been seen from west of the Rockies.
In Florida it has been found in nearly all areas
except the Everglades and the Keys (with the exception
of a doubtful record from Key West by Blatchley,
1928). The original description mentioned only "Amer-
ica," but Robinson (1948b:93) listed the type locality
as "New York."

BIOLOGY: As the common name "tumble bug" im-
plies, these beetles roll balls of dung. The adults and
larvae feed primarily on horse and cow dung, although
they have been found on a variety of other types.
They are not normally necrophagous, but Bragg (1957)
reported a situation where dead tadpoles of Scaphiopus
holbrooki hurteri were used in constructing the balls.
Since this presumably occurred in Oklahoma, this rec-
ord could refer to either pilularius or imitator. The
life history was described by Lindquist (1935), Cooper
(1938a), Ritcher (1945), and Miller (1954). The be-
havior has only recently been studied in detail by
Matthews (1963). From this study it was determined
that the brood balls are rolled solely by the male, with
the female often riding atop but not participating in
the rolling operation. Previously it had often been
reported that the sexes cooperate in this chore. Mat-
thews also discovered that there may be some sun
orientation in rolling the balls. Further study in this
area would be very interesting. Two kinds of balls are
formed; one for adult food and the other for pro.
visioning the larva. For a general discussion of ball
rolling in dung beetles, see the discussion on nidifica.
tion in Halffter and Matthews (1966).
The larval epipharynx was first described by Hayes
(1929), but the figure is based on a broken specimen,
as pointed out by Ritcher (1945:4). The third instar
larva was subsequently described and figured by
Ritcher (1945:6-7).
The larva of none of our other species has been
described for comparison. It can be separated from

the other known larvae of the subfamily Scarabaeinae
by the following combination of characters: prothoracic
shield with an anteriorly projecting, angular process
on each side; legs with a single terminal seta; venter of
last abdominal segment with a single, broad, caudal,
median lobe; median portion of venter of last abdomi-
nal segment with two inconspicuous patches of very
short setae.
Although there is probably more published infor-
mation on this species than any other dung beetle,
much is yet to be learned about behavior and morpho-
logical variation:

MORPHOLOGY: The internal anatomy was studied by
Cooper (1938b), and it is this study which is the
basis for most generalizations on dung beetles. The
abdominal ganglia of the nervous system are coalesced
with the meso- and metathoracic ganglia to form a
single ganglionic center. The stomodaeum has neither
crop nor proventriculus. The mesenteron is exception-
ally long, with projecting gastric caeca for its entire
length. All body cavities are lined with sausage-
shaped fat bodies formed around tracheal branches.
The reproductive system of the female has only the
left ovary developed, with a single ovariole. The res-
piratory system shows no air sacs arising from ab-
dominal tracheae, but seven pairs arise from the
metathoracic spiracles and form a group caudad to
the heavily muscled thoracic region. Air sacs from the
mesothoracic spiracles extend forward into the head.
The external morphology was described by Mohr
(1930) and Halffter (1961). The mouthparts were de-
scribed in detail by Hardenburg (1907) and Miller

SPECIMENS EXAMINED: Over 600, of which 332
were from Florida (for complete data see Appendix 2).

SELECTED REFERENCES: The more important ref-
erences are cited in the above discussions. Since there
are over 100 citations for this species, no attempt is
made to list them here, but the reader is referred to
the catalogue of Vulcano and Pereira (1964) for a
nearly complete listing.

Canthon vigilans LeConte
(Fig. 85, 87)
Canthon vigilans LeConte 1858:16.
DIAGNOSIS: Distinguished from the two other Florida
species by the greater width of the eye as seen dorsally
(Fig. 85). It differs further from chalcites by having
the head, pronotum, and pygidium with coarse granules
as on the elytra. The color is fairly uniform black
with very slight blue to purple reflections; never
bronze or green as in the other species. It is exclu-
sively nocturnal. It reaches the maximum size for the
genus (length 22 mm).

TAXONOMIC NOTES: Halffter (1961) indicated that
this was the least variable of the North American

Canthon, but he showed (Fig. 40 and 41) north-south
clinal variation in two mouthpart characters. Addi-
tional specimens will be required to determine the full
extent of this variation.

Originally described from Georgia, Missouri, and Texas,
without specific designation of a type locality. Robin-
son (1948a:95) listed the type locality as "Texas,"
and he was followed by Halffter (1961). It has been
recorded from the following states: Alabama, Arkansas,
Delaware, District of Columbia, Florida, Georgia, Illi-
nois, Kansas, Massachusetts, Michigan, Missouri, Mis-
sissippi, New Jersey, New York, North Carolina, Texas,
and Virginia. I have also seen specimens from Ken-
tucky and Tennessee (new state records).
It was first reported from Florida (Ft. Myers) by
Blatchley (1928a:68-69). I have not seen specimens
from that part of the state, my southernmost record
being Pasco County. It is probably more widely dis-
tributed than the few records indicate.

BIOLOGY: Practically nothing is known about this
species except that it is nocturnal. It has been taken
from March through September in Florida, nearly al-
ways at light. The two other Florida species are not
attracted to light. Brown (1928a:25) recorded it from
Oklahoma in excrement of horses and cattle on soil
that contained little or no sand. Obviously such a
situation does not exist in Florida, indicating a range
of edaphic tolerances. Miller (1954:Table 1-2) found
this species attracted to traps containing human feces
at night in Georgia. The immature stages are unknown.

40 0.

SPECIMENS EXAMINED: About 125, of which 84
were from 18 Florida localities (for complete data see
Appendix 3).

SELECTED REFERENCES: Blanchard, 1885:166;
Blatchley, 1928a:68-69; 1928:61; Brown, 1928a:25;
Halffter, 1961:294-297, Fig. 34, 68-70, 17; Horn, 1870a:
47; Robinson, 1948b:94-95, Fig. 4; Vulcano and
Pereira, 1964:635-636.

(Fig. 58, 100, 106)

Boreocanthon Halffter 1958:208-209.

TYPE SPECIES: Canthon ebenus (Say), by original

DIAGNOSIS: Similar in general appearance to Can-
thon. Dull gray, matte, granular, clypeus with four
teeth (the genal angles sometimes resembling two
more teeth). The main character for separating this
genus from Canthon (sensus strictus) is the absence
of a marginal line on the anterior part of the posterior
femur (Fig. 106). It is easily separated from Melano-
canthon, which it closely resembles superficially, by the
single spur on the posterior tibia (Fig. 58).

TAXONOMIC NOTES: Halffter (1958) indicated that
he was only describing the genus, and no attempt was
made at a generic revision. Vulcano and Pereira (1964)
listed 12 species in the genus, and Howden (1966b)
added another. Many of the species are variable, and
they are not well known. The genitalia are sometimes
useful in distinguishing closely related species. The
two Florida species are easily separated by the char-
acters given in the key.

composed of nine species, appears to be exclusively
North American. The distribution given by Halffter
(1958:208) is as follows: British Columbia to Alberta
in Canada; United States; northwest of Mexico (Baja
California, Sonora, and Chihuahua).

BIOLOGY: Very little has been published on the
habits of the species except that they are coprophagus.
Presumably they roll balls of dung, as does the related
Canthon, but there are no published records to this
effect. At least one of our species (probus) appears to
be more abundant on rabbit pellets. The immature
stages are unknown.

SELECTED REFERENCES: Halffter, 1958:208-210,
1961:234; Howden, 1966b:729-730; Robinson, 1948b;
Vulcano and Pereira, 1964:595-600.

Key to the Florida species of Boreocanthon
1. Head, pronotum, elytra, and pygidium with en-
larged granules scattered over the minutely peb-
bled surface; anterior tibial spur of the male
elongate, pointed, not bifurcate; larger (length
7-10 mm) ............... depressipennis (Lec.)
1'. Head, pronotum, elytra, and pygidium without
enlarged granules on the minutely pebbled surface,
instead they are replaced by small shining spots;
anterior tibial spur of the male bifurcate; smaller
(length 4-6 mm) ...............probus (Germ.)

i i.

Fig. 100. Boreocanthon depressipennis (Lec.), line = 5mm.

Boreocanthon depressipennis (LeConte)
(Fig. 100, 106)
Coprobius depressipennis Dejean 1836:152 (nomen
Canthon depressipennis LeConte 1859a:11.
Canthon depressipenne Lec., Blackwelder 1944:199.
Boreocanthon depressipennis (Lec.), Halffter 1958:208.
DIAGNOSIS: Easily distinguished from our only other
species by the characters in the key. In addition speci-
mens sometimes have green or blue reflections. The
male genitalia is also distinctive. It is most similar to
B. ebenus (Say) of the southwest, but differs by the
less coarsely granulate pronotum, external elytral striae
less deep than those of the disc, posterior femur with
scattered punctures and fine setae, and the shape of
the male genitalia.

TAXONOMIC NOTES: A readily recognized species,
with no taxonomic confusion or synonyms.

Originally described from Kansas, although Robinson
(1948b:92) erroneously listed the type locality as
Georgia. It has been recorded from the following states:
Louisiana (Summers, 1874:87); "Da., Fla., Ga., Kans."
(Blanchard, 1885:165); Ohio (Dury, 1902:153); North
Carolina (Brimley, 1938:199); Alabama (Loding, 1945:
98); South Carolina (Robinson, 1948b:92); Gillet
(1911:92) erroneously listed it from "Sudamerika." It
is also listed from Mexico by Blackwelder (1944:199)
and Vulcano and Pereira (1964:596), although I have
been unable to verify these records. I suspect that
they refer to B. ebenus.
In Florida it was first recorded from Tampa as
rare by Schwarz (1878:449). Slosson (1893:150) re-
ported it from Suwannee Springs, and Castle and Lau-
rent (1896:303) listed it from Enterprise. Dozier (1918
and 1920) listed it as common at Gainesville. Blatchley
(1928:61) reported it "Throughout the State" and
mentioned a manuscript record by Schwarz from Key
West. I have been unable to verify the latter record
by specimens, and I doubt its validity, since I have
seen no specimens from south of St. Petersburg. It
probably occurs throughout the panhandle and as far
south as the center of the peninsula.
BIOLOGY: A fairly common species in cow dung in
open sandy pastures. Dozier (1918:332) mentioned that
it was common around dung in roads from April to
September. My records are for February through Sep-
tember. Nothing else seems to have been published on
its habits, and the immature stages are unknown.

SPECIMENS EXAMINED: 224 from 24 Florida locali-
ties (for complete data see Appendix 4).

SELECTED REFERENCES: Blanchard, 1885:164-165;
Blatchley, 1928:61; Halffter, 1958:208-209, Fig. 2;
Horn, 1870:46; Vulcano and Pereira, 1964:596.

Boreocanthon probus (Germar)

Ateuchus probus Germar 1824:98.
Canthon minor Sturm 1843:104.
Canthon probus (Germ.), LeConte 1863b:36.
Canthon probum (Germ.), Blackwelder 1944:201.
Boreocanthon probus (Germ.), Halffter 1958:208.

DIAGNOSIS: Easily distinguished from our only other
species by the characters in the key. In addition to
the large granules being replaced by shining spots, the
head and pronotum are minutely, very shallowly punc-
tate (often only visible at an oblique view at high
[90X] magnification). It is superficially similar to
melanus Robinson, but the pronotal punctures are
smaller, shallower, and more widely scattered, and
the male genitalia are different.

TAXONOMIC NOTES: Horn (1870:45) synonymized
abrasus, stating "I have no hesitation in uniting the
species of LeConte to that of Germar, and although
some slight differences exist between the description
and LeConte's unique, it must be remembered that
both species have been described from single speci-
There is some variation in size (length 4-6 mm)

and in the coarseness of the pebbled surface of the
head, pronotum, and elytra. The single specimen I have
seen from Texas has the pebbles more noticeable and
in greater relief, although I can find no further dif-
104). It was originally described from "America septen-
trionali," and Kansas was the type locality for the syno-
nym abrasus. In addition it has been recorded from
the following states: Florida, Georgia, Kentucky, South
Carolina (Blanchard, 1885:165); New Jersey (Smith,
1910:313); North Carolina (Brimley, 1938:199); Ala-
bama (Loding, 1945:98); Oklahoma, Texas, Utah, and
Virginia (Robinson, 1948b:90). I have also seen
specimens from Arizona (new state record).
It was first recorded from Florida (Enterprise) by
Schwarz (1878:449). Blatchley (1918:54) added Cres-
cent City and St. Augustine, and later (1928:61) in-
cluded Centerville, St. Mary, Marion Co., and Dunedin.
My records include nearly the entire peninsula, but
there are no records west of Leon Co. However, this
is probably an artifact of collecting, since it was re-
corded from Mobile Co., Alabama (Loding, 1945).
BIOLOGY: I have found this species to be rather un-
common in Florida. However, this observation is based
on collecting in cow dung in pastures. It appears to be
more abundant in sandy, wooded situations. Several
specimens were taken at Gainesville in rabbit pellets,
and Miller (1954:Table 1) reported 277 specimens
from can traps using human feces as bait. Of this total,
28 were taken in the daytime, and 249 were taken at
night. There is a single specimen labeled "at light,"
but I suspect this is an incidental record. I have taken
specimens on two occasions in malt bait traps, but
this does not appear to be a good attractant. It has
been taken in Florida every month except September
and December. The immature stages are unknown.
SPECIMENS EXAMINED: 61, of which 48 were from
22 Florida localities (for complete data see Appen-
dix 5).

Fig. 105-106. Ventral view of left posterior femur: 105)
Canthon pilularius (L.), note marginal line at arrows. 106)
Boreocanthon depressipennis (Lec.), note lack of marginal line.

SELECTED REFERENCES: Blanchard, 1885:165;
Blatchley, 1918:54, 1928:61; Halffter, 1958:210, Fig.
3; Horn, 1870:45; Miller, 1954:380-381, Table 1-2;
Robinson, 1948b:89-90, Fig. 19-20; Vulcano and
Pereira, 1964:598-599.

(Fig. 57, 107)
Melanocanthon Halffter 1958:210-211.

TYPE SPECIES: Canthon bispinatus Robinson,
1941:128 (by original designation of Halffter
DIAGNOSIS: Superficially similar to Canthon and
Boreocanthon, but differing from both, and from all
other members of the subfamily Scarabaeinae, in pos-
sessing two spurs on the posterior tibiae. Clypeus quad-
ridentate, the genal angles often acute and resembling
a third pair of teeth. Medium sized (length 6-10 mm),
dull, matte, gray to black, dorsal surface granular.
Setae separating the submentum from the gula forming
a V-shaped line, elongate posteriorly at the middle.
Posterior femur without anterior marginal line. Elytral
striae obsolete. Pygidium with a basal transverse carina.
TAXONOMIC NOTES: Most of the species are not well
known, and the genus needs revision. In many cases
the male genitalia are the only reliable characters for
identification. Only four species were listed by Halffter
(1958) and Vulcano and Pereira (1964).

confined to the U. S. east of the Rocky Mountains,
from New Jersey to Florida and Texas to Nebraska.
Three of the four species are found in Florida, the
remaining one (nigricornis Say) is essentially a Great

I . ..

Fig. 108. Stereoscan photo (30X) of Melanocanthon granuli-
fer (Schm.) head (caudal view).

Fig. 109. Stereoscan photo (244X) of right eye of Melano-
canthon granulifer (Schm.) (dorsal view). Note carina sur-
rounding eye and granules of head enlarged in Fig. 110.

Fig. 107. Melanocanthon punctaticollis (Schffr.), line = 5mm.

Fig. 110. Stereoscan photo (1090X) of head granules of
Melanocanthon granulifer (Schm.).

Plains species.
BIOLOGY: The species usually are rarely collected.
For instance, when Robinson (1941) revised the group
he saw only 86 specimens in the 5 collections studied
(including USNM, ANSP, and AMNH). He mentioned
that most of his specimens of bispinatus were taken
in the autumn, usually on old, partly dried toadstools
in the pine barrens of New Jersey. At least some of
the species feed also on cow dung, and I have collected
all three of the Florida species in malt bait traps.
SELECTED REFERENCES: Halffter, 1958:210-212,
Fig. 7-11; Robinson, 1941:127-130, Fig. 1-8; Vulcano
and Pereira, 1964:594-595.

Key to the Florida species of Melanocanthon
1. Pronotal granules reduced to shining spots in
central area, none in relief; head and pronotum
noticeably punctate; Fig. 119-121 ..........
................. punctaticollis (Schaeffer)
1'. Pronotal granules in relief (Fig. 108), although
sometimes reduced in the antero-median area;
pronotum not noticeably punctate .........2
2(1'). Granules of dorsal surface more dense, in
greater relief; punctures of head barely notice-
able; male genitalia as in Fig. 113. ..........
..................... granulifer (Schmidt)
2'. Granules of dorsal surface less dense, less
raised; punctures of the head as noticeable and
as evenly distributed as the granules; male
genitalia as in Fig. 114.. bispinatus (Robinson)

Melanocanthon bispinatus (Robinson)
(Fig. 114)
Canthon bispinatus Robinson 1941:128-129, Fig. 2-3.
Melanocanthon bispinatus (Robinson), Halffter 1958:
DIAGNOSIS: Although the male genitalia (Fig. 114)
are very distinctive, this species is otherwise often diffi-
cult to separate from granulifer without comparative
material. In general the granules are less pronounced
and often reduced to elongate, shining, black spots in
the antero-median area of the pronotum. The punctures
of head and pronotum, although perhaps no more nu-
merous, are more noticeable because of the greater
space between the granules.
TAXONOMIC NOTES: A certain amount of variation
occurs in the extent of granulation and punctures, but
the male genitalia seem to be constant. I can see no
differences between the genitalia of Florida specimens
when compared to Robinson's figure of the type from
New Jersey. Some of the older records of nigricornis
(Say) probably refer to this species. Although origin-
ally described in Canthon, bispinatus was made the
type of the genus Melanocanthon by Halffter (1958:
The type locality is Warren Grove,* Burlington Co.,

New Jersey. Paratypes were recorded from the follow-
ing states: Alabama, Florida, Georgia, New Jersey,
North Carolina, Rhode Island, South Carolina, and
In Florida it has been reported from DeFuniak
Springs and Enterprise (Robinson, 1941:129). My
records add eight additional localities, all in the
northern one-third of the peninsula. It is possible that
part of the records for nigricornis of Blatchley (1928:
60-61) refer to this species or to granulifer. M. nigri-
cornis is known from Texas to Michigan and Nebraska,
but it is not definitely known from the southeast.
BIOLOGY: The only published note on this species is
by Robinson (1941:127) who stated: "I have collected
most of my specimens of bispinatus in the autumn,
usually on old, partly dried toadstools in the pine
barrens of New Jersey. Only once have I seen a speci-
men rolling a ball and this particular ball was composed
of deer excrement." My Florida specimens were taken
primarily in malt bait traps and unbaited pitfall traps.
Two specimens were taken by Wilson Baker on a dead
roach on a mammal snap trap. Specimens have been
taken from February through September, but in Florida
they seem to be most abundant in April. The immature
stages are unknown.
SPECIMENS EXAMINED: 122, of which 106 were
from 11 Florida localities, representing 74 collection
records (for complete data see Appendix 6).
SELECTED REFERENCES: Blatchley, 1928:60-61 (as
nigricornis?); Halffter, 1958:211-212, Fig. 11; Vulcano
and Pereira, 1964:594.

.,r 'g

Melanocanthon granulifer (Schmidt)
(Fig. 108-10, 113, 115-16)

Canthon granulifer Schmidt 1921:126.
Melanocanthon granulifer (Schmidt), Halffter 1958:211.
Melanocanthon granulifera (Schmidt), Howden 1966b:
730 (misspelling).

DIAGNOSIS: Although the male genitalia (Fig. 113)
are very distinctive, this species is otherwise often diffi-
cult to separate from bispinatus without comparative
material. In general the granules are everywhere more
dense and with greater relief. Antero-median area of
the pronotum densely granulate as the remainder, never
with any extensive areas where the granules are re-
duced to shining spots. Punctures of head and pronotum,
although perhaps no less numerous, not as noticeable
because of the density of granules.

TAXONOMIC NOTES: I have seen only a few speci-
mens from Texas (type locality), and they seem to
differ slightly from those from Florida. Further study
of additional material will be necessary to determine
if the two disjunct populations are distinct. At least
some of the early records for nigricornis refer to this

It has been recorded only from Texas and Florida
(Robinson, 1948b:87). At least part of Blatchley's
records of nigricornis (1928:61) probably are refer-
able to this species. My records for Florida include
Manatee Co. in the south to Franklin Co. in the north.
Howden (1966b:730) recorded a specimen from "Mex-
ico" without exact locality.

BIOLOGY: Robinson (1948b:87) collected several
specimens of this species in a cow pasture on April 15,
at Romeo, Florida, where they were all rolling balls

113 114

Fig. 113-114. Caudal view of male genitalia: 113) Melano-
canthon granulifer (Schm.), 114)M. bispinatus (Rob.).


of dung over the ground. Miller (1954:380, Table 1)
reported nine specimens in the daytime and five at
night from traps containing human feces. I have taken
specimens in malt bait traps, in fungi, cow dung, on
dead bird, and on dead cottonmouth. Other habitat
data includes under rotten citron and in Japanese
beetle trap. In Florida it probably occurs throughout
the year, although records are lacking for November
and December.
SPECIMENS EXAMINED: Three from Texas and 129
from 18 Florida localities, representing 34 collection
records (for complete data see Appendix 7).
SELECTED REFERENCES: Balthasar, 1939:180;
Blatchley, 1928:61 (as nigricornis?); Halffter, 1958:
211-212, Fig. 9; Howden, 1966b:730; Miller, 1954:380,
Table 1, Fig. 5; Robinson, 1941:130, Fig. 7-8; 1948b:
87, Fig. 43-44; Schmidt, 1922b:61, 75; Vulcano and
Pereira, 1964:594.

Melanocanthon punctaticollis (Schaeffer)
(Fig. 107, 117-21)
Canthon nigricornis var. punctaticollis Schaeffer 1915:
Canthon punctaticollis Schffr., Blatchley 1928:61.
Melanocanthon punctaticollis (Schffr.), Halffter 1958:
DIAGNOSIS: Easily distinguished from the other three
species of the genus by the reduction of the pronotal
granules to bare, shining spots and the dense puncta-
tion. The male genitalia are also distinctive.

Fig. 119-121. Stereoscan photos of pronotal granules of
Melanocanthon punctaticollis (Schffr.): 119) 85X, 120) 180X,
121) 855X. *

TAXONOMIC NOTES: Although it was originally de-
scribed as a variety of nigricornis, it is a distinct
species, as pointed out by Blatchley (1928:61) and
Robinson (1941:128).
It is apparently confined to north-central Florida from
High Springs in the north to Arcadia in the south.
BIOLOGY: Nothing has been published on the habits
of this species, except the record by Blatchley (1928a:
68) of a single specimen beneath cow dung. I have
taken it in fair numbers in malt traps set in turkey
oak scrub, where large numbers of Peltotrupes pro-
fundus and Mycotrupes gaigei were found. Specimens
have been taken from February through July. The
immature stages are unknown.
SPECIMENS EXAMINED: 270 from 14 Florida local-
ities representing 24 collection records (for complete
data see Appendix 8).
SELECTED REFERENCES: Blatchley, 1928:61; 1928a:
68; Halffter, 1958:211-212; Robinson, 1941:128, Fig.
1, 2; 1948b:86, Fig. 41-42; Vulcano and Pereira, 1964:

(Fig. 122-24)
Glaphyrocanthon Martinez 1948:41.
TYPE SPECIES: Glaphyrocanthon variabilis Martinez
1948 (by original designation).
DIAGNOSIS: Characterized in our fauna by the small
size (length 2-4 mm), usually bright, shining metallic


Fig. 122. Glaphyrocanthon v. viridis (Beauv.), line = 3mm.

green color, bidentate clypeus, and more oval body
shape. The smallest member of the tribe in Florida,
but approached in size by Pseudocanthon perplexus
Lec. and Boreocanthon probus (Germ.). Distinguished
from both genera by the bidentate rather than quad-
ridentate clypeus.
TAXONOMIC NOTES: The genus was divided into
two subgenera (Glaphyrocanthon and Coprocanthon)
by Martinez (1950:160). Coprocanthon contains four
species, none of which extends to North America. The
nominate subgenus is the larger of the two and con-
tains our only U. S. species. The most recent treatise
on the genus is that by Martinez, Halffter, and Halffter
(1964), in which 28 species are included. Our single
species is divided into seven subspecies. Howden
(1966b) added a new species from Mexico, to bring
the known species to 29.
primarily Neotropical, with only a single species ex-
tending north into the U. S., and only one other
species is known from Mexico. The genus is absent
from the West Indies.
BIOLOGY: The species are nearly all confined to
forests (selva) where they have a variety of feeding
habits. Some species are associated with the dung of
tapirs and monkeys. G. viridis leechi has been found
in a hymenomycetous fungus (Halffter, 1959:170).
Most of the species are diurnal, although there are a
few records of G. v. viridis at light. Specimens of
some species can be collected with traps using
rotten meat or malt with proprionic acid. The immature
stages are unknown for all the species.
SELECTED REFERENCES: Halffter, 1961:232; How-
den, 1966b:728-729, Fig. 1-2; Martinez, Halffter, and
Halffter, 1964:1-42, Fig. 25; Pereira and Martinez,
1956:125-134; Vulcano and Pereira, 1964.:660-666.

Glaphyrocanthon v. viridis (Beauvois)
(Fig. 122)
Copris viridis Palisot de Beauvois 1805:24, P1. 3, Fig. 2.
Ateuchus obsoletus Say 1823:208.
Onthophagus viridicatus Say 1835:173.
Canthon viridulus Dejean 1836:152 (nomen nudum).
Canthon metallicus Sturm 1843:104.
Canthon viridis Beauv., LeConte 1859b:11.
Canthon viride Beauv., Blackwelder 1944:202.
Glaphyrocanthon viridis (Beauv.), Pereira and Martinez
Glaphyrocanthon v. viridis (Beauv.), Martinez, Halffter,
and Halffter 1964:21-24, Fig. 3-5.
DIAGNOSIS: Small (length 2-4 mm), bright metallic
green or copper colored, shining, the clypeus bi-
dentate. Our only representative of the genus, which
can be readily distinguished by the characters in the
TAXONOMIC NOTES: The species is extremely vari-
iable, and seven subspecies were created by Martinez,
Halffter, and Halffter (1964). However, Howden
(1966b:729) suggested that some of these are full
species, but more material is needed before the prob-
lems can be resolved. Typical viridis is the only sub-
species reported from the U. S. It is quite variable
over its broad range here, and perhaps other subspecies
could be described. Large series of specimens are not
available and, until they are, the variation cannot be
properly evaluated. The color is normally a shiny
green, but occasionally specimens are coppery and less


shiny. All such specimens that I have seen are from
northern localities.

24). The type locality of this subspecies is "caroline
du Sud des Etats-Unis d'Amerique." It is possible
that this refers to either North or South Carolina. It
has been recorded from nearly all of the eastern U. S.
from New York to Florida, and from Minnesota to
Monterrey in Mexico.
The Florida distribution is spotty, with nearly all
records being from the north central and eastern por-
tion. The two records from the southern half of the
state are based on the literature; Lake Worth (Blatch-
ley, 1928:61) and Miami (Martinez, Halffter, and
Halffter, 1964:24). The species is seldom collected, and
the few records probably do not reflect the true dis-

BIOLOGY: This is one of the few Florida dung beetles
which seems to be confined to densely wooded areas
(hammocks). Although Martinez, Halffter, and Halffter
(1964:11) reported collecting specimens at light at
Monterrey and Apodaca, Mexico, I have never taken it
in over 1000 Florida light trap samples examined.
Blatchley (1910:914-915) found it scarce in Indiana
". .. beneath flat stones on hillsides near Wyandotte
Cave." Three specimens were taken in malt traps to
which a few drops of proprionic acid were added. I
collected a single specimen rolling a pellet of rabbit
dung along a path in dense woods near Citico, Ten-
nessee. This was a mountainous area where a good
series of Ataenius brevis Fall was taken along the
same path. Halffter and Matthews (1966:21) reported
an unpublished record by Cartwright of this species
being found on leaves around bird droppings. The
greatest number of Florida specimens was taken in
unbaited pitfalls at Tall Timbers Research Station
(Leon Co.). In Florida it has been collected from
March through September, with a single record for
December. Cartwright (1934b:238) recorded it in South
Carolina from February through August. The immature
stages are unknown.
SPECIMENS EXAMINED: 280, of which 223 were
from 8 Florida localities, representing 96 collection
records (for complete data see Appendix 9).
SELECTED REFERENCES: Bates, 1887:30-31; Blan-
chard, 1885:167; Blatchley, 1910:914-915; Dillon and
Dillon, 1961:509, P1. 49, Fig. 1; Halffter, 1961:246;
Horn, 1870a:47; Robinson, 1948b:97, Fig. 15-16; Vul-
cano and Pereira, 1964:665-666.

(Fig. 125-127)
Pseudocanthon Bates 1887:35.
Opiocanthon Paulian 1947:30.
TYPE SPECIES: Canthon perplexus Lec., 1847 (by
DIAGNOSIS: Small (length 3.5-5 *mm), oval, black,

sometimes with slight green or purple reflections,
shining. Clypeus with four reflexed teeth, the middle
two larger; slight angulation at the geno-clypeal suture.
Eyes prominent, elongate dorsally. Pronotum appearing
subparallel-sided from above, the sides margined, but
the base without marginal line. Anterior part of pro-
thorax excavated beneath to receive antennal club and
anterior tibia, the excavation bounded posteriorly by a
transverse margin. Mesosternum elongate. Elytra with
nine weakly impressed striae. Pygidium without basal
transverse carina (in our single species). Dorsal surface
minutely alutaceous (or pebbled) but not roughly gran-
ular as in Canthon, Boreocanthon, and Melanocanthon.
Only two other Florida members of the tribe
(Glaphyrocanthon viridis and Boreocanthon probus)
approach the small size of this species. From the first
it can be distinguished by the quadridentate (Fig. 125)
rather than bidentate (Fig. 122) clypeus; from the latter
it differs by having only a slight angle at the geno-
clypeal suture, by the lack of a basal transverse carina
on the pygidium, and by the presence of a posterior
carina bordering the excavated area of the prothorax.


Fig. 125. Pseudocanthon perplexus (Lec.), line = 3mm.

TAXONOMIC NOTES: Although this genus was de-
scribed long ago, it was supressed under Canthon by
most authors until recently. It was recognized as valid
by Arrow (1903), but not amplified until Martinez
(1947a). Although several authors doubt the validity
of other recently described genera of this tribe, they
are all in agreement that Pseudocanthon merits generic
status. Matthews (1966:85) stated that "In spite of
the disagreement among specialists as to which fea-
tures are of generic importance, there can be no doubt
that Pseudocanthon is a valid genus, with an un-
mistakable facies." He synonymized Opiocanthon
Paulian in the same paper.

The males have the spur of the anterior tibia bi-
furcate, elongate on the outside and rounded on the
inside; the female has this spur elongate and curved,
but not bifurcate.
The genus is not well known and, as evidenced
by several new species recently being found in the
West Indies, there will probably be others discovered.

are listed by Vulcano and Pereira (1964), one of which
(P. perplexus) occurs in the U. S., Central America,
Colombia, Venezuela, and Brazil. Of the other two,
chlorizans Bates is found in Mexico, the West Indies,
and Colombia. The remaining species, xanthurus Blan-
chard, is recorded from Colombia, Brazil, and Argen-
tina. Matthews (1966) described four new species from
the West Indies and transferred another West Indian
species from Opiocanthon to Pseudocanthon. Thus, there
are now eight species in the genus.

BIOLOGY: All of the species apparently are copropha-
gous, some species (not perplexus) being common in
cow and human dung. Our single species is rarely
collected except at light, but has been taken in packrat
(Neotoma sp.) nests, carrion, and rotting cantaloupe.
For additional information see this section under per-

SELECTED REFERENCES: Bates, 1889:386; Halffter,
1961:232-233; Martinez, 1947a:263-267; Matthews,
1966:83-99; Pereira and Martinez, 1956:109; Vulcano
and Pereira, 1964:591-592.

Pseudocanthon perplexus (LeConte)
(Fig. 125)
Canthon perplexus LeConte 1847:85.
Pseudocanthon perplexus (Lec.), Bates 1887:35.
Canthon perplexum Lec., Blackwelder 1944:200.

DIAGNOSIS: Readily distinguished from the other
Florida members of the tribe Scarabaeini by the
characters in the key and those listed under the gen-
eric diagnosis. It can be separated from all the West
Indian species, except chlorizans Bates, by the lack of
a basal transverse carina on the pygidium.

TAXONOMIC NOTES: This species is variable in color,
from black with practically no metallic reflections to
extensive green or purple reflections. Variation is also
notable in the punctation of the pronotum and elytral
intervals, and in the degree of alutaceousness. In some
specimens the elytral intervals are completely aluta-
ceous and almost granulate, and in others there is
hardly a trace of this sculpture. It is very similar to
chlorizans Bates, and, as Howden (1966b:733) sug-
gested, the two may represent geographical races of
a single species. I have examined a specimen de-
termined as chlorizans by Bates, and I am unable to
separate it from examples of perplexus from Texas.
Further study of a large series of specimens, from all
parts of the range, will be necessary before the exact
status of this species can be determined.

dO .''

It was originally described from Quincy, Illinois, al-
though Robinson (1948b:97) erroneously listed the type
locality as Texas. It was recorded by Vulcano and
Pereira (1964:592) from ". . . North and Central
America, Colombia, Venezuela, and Brazil (Amazonian
region)." Published records have been found for the
following states: Alabama (Loding, 1933:147); Ar-
kansas, California, Illinois, Texas (Blanchard, 1885:
167); Florida (Blatchley, 1918:54); Indiana (Blatchley,
1910:915); Louisiana, Arizona (Robinson, 1948b:97).
In addition to these states, I have seen specimens from
Kentucky, Mississippi and Missouri (new state records).
The record of "Cal." by Blanchard (1885:167) is
questionable, even if it refers to Baja California.
In Florida it has been previously reported from
Dunedin, Royal Palm Park, and Enterprise (Blatchley,
1928:62). My records include nearly the entire state
as far south as Grassy Key (Monroe Co.).
BIOLOGY: Although a fair number of specimens was
examined (85), the species is not often collected. The
greatest number collected at one time was four in a
light trap. Very little is known about the biology,
since nearly all specimens have been taken at light.
Blatchley (1928:62) reported taking several specimens
in ". .. bottle bait of amyl acetate and molasses." I
collected a dead specimen on the carcass of a cow and
four specimens in a dung chamber in the nest of
packrats (Neotoma floridana small Sherman) on Key
Largo, Fla. I have seen a single specimen collected on
rotting cantaloupe. From these limited observations it
appears that this species has a wide variety of foods,

but it has not been found in human or cow dung. It
has been taken on eight occasions in unbaited pitfalls
at Tall Timbers Research Station (Leon Co.). Speci-
mens have been collected in Florida every month except
November, January, and February. The immature stages
are unknown.
SPECIMENS EXAMINED: 85, of which 71 were from
29 Florida localities, representing 48 collection records
(for complete data see Appendix 10).
SELECTED REFERENCES: Bates, 1887:35; Blanchard,
1885:167; Blatchley, 1928:62; Horn, 1870a:46; Robin-
son, 1948b:97, Fig. 17-18; Vulcano and Pereira, 1964:

(Fig. 1, 128, 133, 136, 145)
According to Halffter and Matthews (1966:256-259)
the tribe contains 63 genera in the world, a great many
of which are tropical. Only four genera are known
from the U. S., all of which are found in Florida:
Ateuchus, Phanaeus, Copris, and Dichotomius
(=Pinotus). The tribe is represented in Florida by
10 species.
There have been various attempts to divide the tribe
into subtribes. Halffter and Matthews (1966) listed
four subtribes: Dichotomina (Dichotomius and Ateu-
chus), Phaneina (Phanaeus), Coprina (Copris), and
Ennearabdina. Only the last tribe, containing a single
endemic Argentinean genus, is not represented in
Florida. There is little agreement about the placement
of some genera, and until a thorough study of the
higher categories within the subfamily has been made,
the divisions appear too artificial for consideration
The tribe is characterized by the shining appear-
ance of most species, and the nearly glabrous dorsal
surface. The middle and posterior tibiae are expanded
at the apex, and all tarsi are present. The males of
most species have horns and/or protuberances on head/
or pronotum. Except for Phanaeus, the elytral striae
are distinct, and the intervals are smooth and shining.
The Florida representatives vary from 5 to 29 mm in
length. The third segment of the labial palpi is dis-
In Florida the tribe contains our largest dung
beetle, Dichotomius carolinus (L.); two species of
Ateuchus which superficially resemble members of
the family Histeridae; the genus Phanaeus (Fig. 1)
with beautiful metallic red and green colors, as well
as the greatest male head horn development of any
member of the subfamily; and four species of Copris,
three of which are endemic to the state.
Taxonomically the tribe is fairly well known in
the U. S., with a recent revision of the genus Copris
(Matthews, 1961). The genus Phanaeus is especially
in need of a thorough revision; the status of many
color forms is open to question.
Biologically the group has received more attention

than some others because of the stimulating early
work by the French naturalist Fabre (1918). Nearly
all the species are coprophagous or necrophagous. In
contrast to the Scarabaeini, their legs are not well
adapted for ball rolling, and the larval food usually
is provisioned in a wad packed at the bottom of
a burrow. The details of parental care of our species
are not well known, but at least some species of Copris
exercise some care of the dung wad until after pupa-
tion has taken place.

Genus ATEUCHUS Weber
(Fig. 128)
Ateuchus Weber 1801:10.
Ateuchus Fabricius 1801:54.
Choeridium Serville 1828:356 (often cited as 1825)
TYPE SPECIES: Ateuchus histeroides Weber, 1801 (by
DIAGNOSIS: Small (length 5-7 mm), shining, black
to bronze, oval convex, resembling members of the
family Histeridae. Clypeus bidentate, the teeth sepa-
rated by a broad "V" (these teeth sometimes worn
down). Antennae nine-segmented. Scutellum absent.
Pronotum convex, posterior marginal line absent, punc-
tures variable. Eight elytral striae, the intervals convex,
minutely punctate. Sexual dimorphism not obvious,
the spur of the anterior tibia broad and/or truncate
in the male. Posterior tibia without transverse carinae.
Pygidium with a basal transverse carnia. Easily sepa-
rated from the other Florida members of the tribe
Coprini by the transverse, non prominent anterior coxae,
lack of head or pronotal horns and protuberances, and
the small size (the other species 8-20 mm in length).

i . .

Fig. 128. Aleuchus lecontei (Har.), line = 2.5mm.

TAXONOMIC NOTES: The name Ateuchus was used
almost simultaneously by Weber and Fabricius, both
descriptions appearing in 1801. Chapin (1946:79) dis-
cussed this situation and concluded that Weber's paper
had priority. Fabricius included in his Ateuchus sev-
eral Old World species, most of which are now placed
in the genus Scarabaeus. The name Choeridium was
used in most North American literature before 1946,
when Chapin established the synonymy, although Rob-
inson (1948d:37) continued to use Choeridium without
reference to Chapin's paper or to the problem of
Blackwelder (1944:204-205) listed 70 species for
Latin America, and Robinson (1948d:37) recorded
three species and one subspecies from the U. S.
Matthews (1966) described one new species and listed
one other from the West Indies.
Members of the genus are very similar morphologic-
ally and are not well known at present. Although Rob-
inson (1948d) reviewed the U. S. species, his treatment
is not totally satisfactory. It is nearly impossible to
identify the Mexican, Central American, and South
American species with the existing literature. A generic
revision is badly needed. The male genitalia appear to
offer valuable characters, at least in part of the genus.

imately 75 described species, all but 3 inhabit the
Neotropical region. The Antilles have two species, one
of which is endemic to St. Lucia, and the other is


Fig. 129-130. Lateral view of head
Ateuchus spp.: 129) A. lecontei (Har.),
(Web.). Compare position of posterior

and pronotum of
130) A. histeroides
pronptal angle (at

found also in Mexico and Central America. In the U. S.
the three species and one subspecies are found only
east of the Rocky Mountains. One of these occupies
most of the eastern U. S., one is primarily southeastern,
and one is found in Texas and northern Mexico. Two
species are found in Florida.
BIOLOGY: Most of the species are coprophagous or
necrophagous, although at least two are known to be
myrmecophilous. The Florida species are also some-
times found in fungi. The life histories and habits are
not well known. The larva is known for only a single
species (see characterization under A. histeroides).
SELECTED REFERENCES: Blackwelder, 1944:204-
205; Blanchard, 1885:170; Blatchley, 1910:915; 1928:9;
Brown, 1928a:25; Chapin, 1946:79; Gillet, 1911:52-53;
Horn, 1875:137; Matthews, 1966:44-51; Ritcher, 1945:
7-8 (larva); Robinson, 1948d:37-40.

Key to the Florida species of Ateuchus
1. Posterior angle of lateral pronotal margin formed
below the level of the elytra (Fig. 130); head
punctures coarse, at least anteriorly separated by
less than their diameter; male anterior tibial spur
rounded, that of the female elongate, acute; coarse
pronotal punctures present along nearly the entire
lateral margin; more northern species, rare in
Florida .................... histeroides Weber
1'. Posterior angle of lateral pronotal margin formed
at the level of the elytra (Fig. 129); head punc-
tures fine, usually separated by at least twice their
diameter; male anterior tibial spur truncate, that
of the female broadest at the tip; coarse pronotal
punctures rarely present medially at the lateral
margin; a common Florida species (Fig. 128)....
............................ lecontei (Harold)

Ateuchus h. histeroides Weber
(Fig. 130)
Ateuchus histeroides Weber 1801:37.
Ateuchus capistratus Fabricius 1801:62.
Choeridium histeroides (Weber), LeConte 1863b:36.
Choeridium histeroides histeroides (Weber), Robinson
DIAGNOSIS: Generally larger, bronzer, and more
densely punctate than lecontei. Most specimens can be
separated easily by the shape of the lateral pronotal
margin and the other characters mentioned in the key.
The margin is usually reflexed wider and more con-
vex, but these are relative characters. The anterior
tibial spurs can be worn down, especially in the fe-
male, so this character should be used with caution.
TAXONOMIC NOTES: Robinson (1948d) disting-
uished two subspecies, based primarily on the head
and pronotal punctation. The nominate subspecies has
a fairly wide range and appears to be the one present
in Florida. The other subspecies, punctatus Rob., is
more northern (Mich., Ill., Pa.). Robinson's treatment
is inadequate, and a generic revision will be required

to ellucidate the situation. It is likely, judging from the
variability, that there are several entities masquerading
under this name.

occurs throughout the eastern U. S., although much
more commonly in the northeast. Robinson (1948d:39)
recorded it from New Jersey, North Carolina, Okla-
homa, Pennsylvania, and Texas. I have also seen speci-
mens from Kansas, Kentucky, Indiana, and Ohio.
In Florida, Blatchley (1928:9) reported it from
Enterprise, S. Jacksonville, Lakeland, and Gainesville.
I have not been able to locate any of these specimens,
either in the Blatchley collection at Pudue University
or in any of the collections examined. It is possible
that these represent misidentifications of lecontei. I
have seen specimens only from Torreya St. Pk., Florida
Caverns St. Pk., and Newnan's Lake.

BIOLOGY: Apparently this species has a range of
food, from various kinds of dung to rotting fungi. I
collected about 30 specimens in the burrow of a wood-
chuck and several from packrat droppings in Ohio.
Other label data includes human feces, cat carcass,
malt trap, carrion trap, and at light. Apparently light
is not a good attractant. They are most abundant in
wooded situations.
The adults provision the larva with a dung wad
packed into the bottom of a vertical burrow about 10
to 12 inches beneath cow dung. A single egg is laid
near the upper end. Most activity occurs between April
and September.
The larva has been described by Ritcher (1945:
7-8) and is characterized as follows: prothoracic shield
without anteriorly projecting process; third abdominal
segment without a prominent, conical, dorsal gibbosity;
venter of last abdominal segment with two monostich-
ous, longitudinal palidia; width of head capsule of
third instar 2.0-2.1 mm.
Adult mouthparts were described by Hardenberg
(1907), and their role during ingestion of helminth
eggs was discussed by Miller (1961). Cram and Jones
(1929) reported studies on these beetles as intermediate
hosts for tapeworms (Raillietina cesticillus and Hymen-
olepis carioca) of poultry and game birds.

SPECIMENS EXAMINED: Over 100, of which only 9
were from 3 Florida localities as follows: (4) Alachua
Co., Newnan's Lake, 15-XII-31, T. M. Little (USNM);
(1) Jackson Co., Florida Caverns St. Pk., 7-VII-58,
T. J. Walker, Jr. (FSCA); (4) Liberty Co., Camp Tor-
reya, 9-IX-29, T. H. Hubbell #70 (USNM).

SELECTED REFERENCES: Blanchard, 1885:170;
Blatchley, 1910:915, Fig. 361; 1928:9; Chapin, 1946:
79; Cram and Jones, 1929:49-51; Davis, 1966:213;
Dillon and Dillon, 1961:509, P1. 49, Fig. 2; Edwards,
1949a:143; Hardenberg, 1907:562, P1. 31, Fig. 17;
Harold, 1868b:50-52; 1873:107; Horn, 1875:137; Rit-
cher, 1945:7-8, Fig. 2, 4, 12, 19, 3p0, 37, 43; Robinson,

Ateuchus lecontei (Harold)
(Fig. 128-9)
Choeridium lecontei Harold 1868b:52-53.
Ateuchus lecontei (Har.), Blackwelder and Blackwelder
DIAGNOSIS: Generally smaller, blacker, and less
densely punctate than histeroides. Most specimens can
be separated easily by the shape of the lateral pronotal
margin and the other characters mentioned in the key.
The margin is barely reflexed, narrow, and less convex,
but these are relative characters. The anterior tibial
spurs can be worn down, especially in the female, so
this character should be used with caution.
TAXONOMIC NOTES: As is true with histeroides, this
is a variable species. Further study will be necessary
to determine whether this is a "composite" species and
whether the variability is correlated with geography or
Robinson (1948d:38) reported it from Florida, Georgia,
New Jersey, and South Carolina. The type locality was
listed as "Vereinigten Staaten," without precise locality.
Leng (1920:248) listed it from "Fla., Ariz.," but I
have been unable to substantiate the Arizona record.
If any species of Ateuchus occurs there it is probably
texanus Robinson and not lecontei. It has also been
reported from North Carolina (Brimley, 1938:199) and
Alabama (Loding, 1945:99).
In Florida it was recorded from Ft. Capron and
Enterprise (Schwarz, 1878:449); Ft. Worth (Hamilton,

. -00

1894:252); Gainesville (Dozier, 1918:332); and from
St. Augustine to Key West (Blatchley, 1928:9). My
records include most of the area between Santa Rosa
Co. and Punta Gorda. I have not seen specimens from
the southern one-fourth of the peninsula.
BIOLOGY: Practically nothing has been published on
the habits of this species. It is extremely common in
Florida under a variety of situations. It is most abun-
dant at light and in cow dung.
Additional label data include armadillo carcass, dog
dung, dead fish, dead crabs, fleshly fungus, under oak
leaves, horse dung, chicken feathers, Steiner trap,
McPhail trap, human feces, Jap beetle trap, Persea
americana (presumably rotting fruit), and bait traps
with malt, yeast, and proprionic acid. Unbaited pitfall
traps at Tall Timbers Research Station (Leon Co.)
produced 338 specimens in 94 weekly collections.
Young and Goff (1939:60) reported trapping specimens
in the mouth of a gopher tortoise burrow at Leesburg,
but indicated that they were probably "casual."
SPECIMENS EXAMINED: Over 1,500, of which 1,451
were from 54 Florida localities, representing 276 collec-
tion records (for complete data see Appendix 11).
SELECTED REFERENCES: Blanchard, 1885:170;
Blatchley, 1928:9; Edwards, 1949a:143; Horn, 1875:
137; Miller, 1954:379, 382; 1961:737, Table 1; Robin-
son, 1948d:38-39.

(Fig. 133)
Dichotomius Hope 1838b:321.
Holocephalus Hope 1838b:323 (in part).
Homocopris Burmeister 1842:77 (cited as 1846 by
Martinez, 1951:140, but fide Blackwelder, 1957:995,
the date for volume 3 should be 1842).
Selenocopris Burmeister 1842:77 (in part).
Pinotus Erichson 1847:108.
Brachycopris Haldeman 1848b (often cited as 1846).
TYPE SPECIES: Copris boreus Olivier, 1789 (by ori-
ginal designation).
DIAGNOSIS: Contains the largest North American
species of the subfamily Scarabaeinae (length 20-30
mm). Our single species can be readily distinguished
from the related genus Copris by the seven elytral
striae and the large size.
TAXONOMIC NOTES: The above synonymy was
established by Martinez (1951:140). Prior to that time
our common species was placed in Pinotus in nearly
all the U. S. literature. Even some of the recent litera-
ture (e.g., Dillon and Dillon, 1961:512) continued to
use the name Pinotus. There are only two species
known from the U. S.: carolinus (L.) and colonicus
(Say). The latter has two horns or tubercles on the
head of the male, while the former has a single horn.
The group is in need of study to determine if the speci-
mens from Mexico and Central America are conspecific
with those of the U. S. The genus is listed in the sub-

Fig. 133. Dichotomius carolinus (L.), line = 8mm.

tribe Dichotomina by Halffter and

Matthews (1966:

Neotropical, except for two species entering the U. S.
One of these (colonicus) is known from Arizona to
Mexico; the other carolinuss) is found throughout the
eastern U. S. and reported by Blackwelder (1944) from
British Honduras, Guatemala, Nicaragua, Costa Rica,
and Panama. There is no accurate count of the number
of species, but Blackwelder (1944) listed 141 species.
BIOLOGY: Most of the species are coprophagous, but
a few have necrophilous tendencies. Little information
is available on the habits of any except our single
SELECTED REFERENCES: Martinez, 1951:138-142;
Schaeffer, 1906:256.
Dichotomius carolinus (Linnaeus)
(Fig. 133)
Copris carolina Linnaeus 1767:125.
Copris monacha Dejean 1836:154 (nomen nudum).
Brachycopris carolina (L.), Haldeman 1848b:125.
Pinotus carolinus (L.), Gemminger and Harold 1869:
Pinotus bituberculatus Harold 1869b:127.
Dichotomius carolinus (L.), Martinez 1951:140.
DIAGNOSIS: Large (length 20-30 mm), black, shin-
ing, very convex, bulky. Head contains a horn or
tubercle, nearer the base in the female and nearer
the apex in the male. Pronotum margined with elon-
gate, curved setae; extremely convex on basal one-half,
and abruptly descending to the head on the anterior
one-half. Pronotum with a deep depression on each
side at the carina behind the declevity. Elytra notice-

ably striate, striae usually filled with dirt for part of
their length posteriorly. This area, actually an enlarge-
ment of the striae fitted with minute stiff setae to
which dirt adheres, broader and more noticeable in
the female. Middle and posterior tibiae greatly ex-
panded at the apex, the spurs elongate and flattened
on one side. The general faces (Fig. 133) should
permit easy recognition of this, our largest dung beetle.

TAXONOMIC NOTES: The nearest relative is colonicus
from the western U. S. and Mexico. This species was
originally described as a variety of carolinus, and its
status needs to be reevaluated. In fact these two forms
probably represent a complex of species, for which
additional study will be required. I believe that some
of the specimens from Mexico and Central America
are not conspecific with those of the U. S.

It is found throughout the eastern U. S. from New
York to Florida west to Texas and north to Nebraska.
It has been recorded from British Honduras, Guatemala,
Nicaragua, Costa Rica, and Panama by Bates (1887),
and I have seen a specimen from El Salvador. The dis-
junction of this range, the apparent absence in Mexico
where colonicus is abundant, and a few slight differ-
ences suggest that more than one species is included
in these records.
In Florida it probably occurs throughout the state,
except for the Everglades and the Keys. Blatchley
(1928:9) recorded it from Suwannee'Springs, Lakeland,

Punta Gorda, Lake City, Gainesville, and Dunedin. I
have seen specimens from 19 localities.

BIOLOGY: This is a common species in cow dung
and at light. Apparently male and female cooperate in
supplying the dung food for the larvae. This is nor-
mally packed in a wad at the end of a burrow which
may be branched or not. Their activity is quite obvious
on the surface where large mounds of soil (sometimes
six inches in diameter) mark the burrow entrance.
Lindquist (1933:112-115) studied the burrowing
behavior in detail in Kansas. By estimating the num-
ber of burrows per acre, averaging the amount of soil
excavated, and the amount of dung buried, he con-
cluded that 21 pounds of dung (air dried) were buried
and 126 pounds of soil excavated per acre. Keeping in
mind that this was only during one point in time, and
that beetles are active a great part of the year, there
can be no doubt about the important role being played
in soil mixing and increased fertility. Australia and
New Zealand are currently considering the introduction
of such dung beetles in order to improve their pastures
(Bornemissza, 1960).
This species is found almost exclusively in open
pastures or roadways and does not occur in forested
areas. It seems to be more abundant where there are
sandy soils. In Florida, specimens have been found
during every month except December and January.
Halffter and Matthews (1966) compiled a list of five
species of Macrocheles mites recorded from this beetle.
The larva is a large "hump-backed" white grub
(Fig. 10). I examined one dung wad containing a
third instar larva and the larva was exposed for photo-
graphing. During an extended delay in taking the pic-
tures the larva repaired parts of the wall by adding
regurgitated food material (Fig. 11). This behavior
has been observed in several other species of dung
beetles (Halffter and Matthews, 1966:182). Howden
(1955a) also mentioned a unique case of the larva of
Peltotrupes building a retreat of its own fecal matter.
The larva has been described by Ritcher (1945:
8-9) and is characterized as follows: prothoracic shield
with an anteriorly projecting, angular process on each
side; legs with a pair of terminal setae, claws absent;
glossa with an irregular transverse row of granules
anterior to the hypopharyngeal oncyli; maximum width
of head capsule of third instar 5.36-6.1 mm.

SPECIMENS EXAMINED: 253, of which 93 were from
31 Florida localities, representing 51 collection records
(for complete data see Appendix 12).

SELECTED REFERENCES: Bates, 1887:52-53; Blatch-
ley, 1910:916, Fig. 362; 1928:9; Dillon and Dillon,
1961:512, P1. 49, Fig. 8; Drury, 1770:77, P1. 35, Fig.
2; Hardenberg, 1907:555-561, Pl. 30, Fig. 1-7, Pl. 34,
Fig. 1; Horn, 1870a:42; Lindquist, 1933:112-115;
Miller, 1954:381; 1961:735, 738, Fig. 5, 6, 8; Mohr,
1943:296; Olivier, 1789:134, P1. 12, Fig. 113; Ritcher,
1945:8-9, Fig. 3, 6, 9, 10, 13, 17, 18, 21, 27, 32, 39
(larva); Schaeffer, 1906:256; Smith, 1892:83, P1. 2,
Fig. 3.

Genus COPRIS Muller
(Fig. 136-44)
Copris Geoffroy 1762:87 (rejected by Int. Cor. Zool.
Nom., Opinion 228).
Copris Muller 1764:xi.
TYPE SPECIES: Scarabaeus lunaris Linnaeus, 1758
(by subsequent designation of Curtis, 1832:414).
DIAGNOSIS: Medium sized (length 8-15 mm), elon-
gate, convex, subparallel, shining. Clypeus broad, mar-
gined, and notched in all Florida species except ine-
marginatus. Sexual dimorphism often striking, the males
with elongate horns on the head and protuberances and
excavations on the pronotum. Easily distinguished by
the general facies (Fig. 136). It can be separated from
Dichotomius, its nearest relative in the Florida fauna,
by the eight elytral striae and smaller size.
TAXONOMIC NOTES: The genus was recently revised
for the Western Hemisphere (Matthews, 1961). Several
of the species (e.g., howdeni Matthews and Halffter, and
halffteri Matthews) are known from very few speci-
The name Copris was first proposed by Geoffroy
1762 (not 1764 as stated by Matthews, 1961:2). How-
ever, the International Commission on Zoological No-
menclature (1954) has discarded this work for nomen-
clatural purposes. Muller (1764) apparently made a
valid "indication" but included no species. The Florida
species are very distinct and easily separated by the
characters in the key.
summary is modified from Matthews (1961:4). The
approximately 160 species are distributed as follows

Fig. 136. Copris minutes (Drury), line = -mm.

(with some species counted twice): Ethiopian (77),
Oriental (46), Palearctic (27), Nearctic (16), Neotropi-
cal (8); it is absent in Madagascar and Australia. It
is the most boreal of the genera of Coprini. In the
Western Hemisphere it is represented in the U. S.
east of the 100th meridian, in all of Mexico and U. S.
territory bordering Mexico (except California), and all
of Central America to Panama. A single Central
American and Mexican species extends to the mountains
of Colombia and Ecuador. The genus is absent in the
remainder of South America, the Antilles, and the
Galapagos Islands. There are presently recognized 23
species and five subspecies from the Western Hemi-
sphere. Nine species and one subspecies are known
from the U. S., of which four species are found in
BIOLOGY: The majority of the species are copro-
phagous, feeding on and provisioning the larvae with
dung of higher mammals. One exception is C. gopheri,
which uses the dung of the gopher tortoise, Gopherus
polyphemus Daudin. There is a slight necrophagous
tendency in some species; I have taken a single C.
inemarginatus on the dry leg tendons of a dead horse,
and Blatchley (1918) found it on a dead turtle.
Apparently all species bury the dung beneath
the source and lay a single egg in each dung ovoid.
The details of the accompanying behavior are variable
among the species. However, there does seem to be
parental care of the brood cell in all the species studied.
The female maintains a smooth contour on the outside
of the ovoid, preventing the growth of fungi and mold.
At least some of the species stridulate. For additional
information on nidification see the papers by Matthews
(1961) and Halffter and Matthews (1966).
The larvae have been described for two North
American species (Ritcher, 1945), and they can be
distinguished from other Scarabaeinae by the following
combination of characters: prothoracic shield with an
anterior, angular projection on each side; legs with a
single terminal seta set on a small blunt claw; venter of
last abdominal segment with paired, median, caudal
lobes, or a cleft median lobe.
SELECTED REFERENCES: Blatchley, 1910:916; 1928:
10; Gillet, 1911:71-79; Horn, 1870a:42-44; Matthews,
1961:1-139; Ritcher, 1945:10-12, Fig. 8, 14, 28, 34,
36, 41, 42 (larvae); Schaeffer, 1906:254-256.

Key to the Florida species of Copris
1. Clypeus entire, at most slightly sinuate medially
(Fig. 138) ............... ................
. inemarginatus Blatch.
1'. Clypeus distinctly emarginate, the central notch
prominent (Fig. 139) ...................... 2
2. Lateral pronotal carina absent; lateral pronotal
margin evenly curved; pygidial margin incomplete,
the inner border effaced ventrally; head armed
with a vertical horn in both sexes; coarse head
punctures absent or confined to genal area; smaller
species (length 7.5-11 mm) ................ 3

2'. Lateral pronotal carina present; lateral pronotal
margin sinuate or slightly angulate; pygidial mar-
gin complete; head of both sexes unarmed; head
with coarse punctures throughout; larger species
(length 13-15 mm); rare species known only from
Oneco and Lake Marion, Fla. ................
................howdeni Matthews and Halffter
3. Coarse punctures of head confined to genal area;
coarse pronotal punctures scattered throughout,
including most of the disc; elytral striae crenulately
punctate, the intervals finely but noticeably punc-
tate; forespur linear, rounded at the tip; common
Florida species .............. minutus (Drury)
3'. Coarse punctures absent on head; coarse pronotal
punctures confined to the anterior angles, the disc
appearing impunctate; elytral striae obsoletely
punctate, never appearing crenulate, the intervals
finely and noticeably punctate; forespur crescent-
shaped, curving outward; confined to burrows of
the gopher tortoise ............ gopheri Hubbard

line_ 2.5mm

partially indicated anteriorly in minutus; completely
absent in gopheri.
TAXONOMIC NOTES: A very distinct and easily
recognized species, but evidently related to minutus.
Matthews (1961:56) placed these two species in his
minutess complex."
Its range is probably coextensive with that of the gopher
tortoise (Fig. 117), but it has not been reported from
outside of Florida. The type locality is Crescent City,
Fla. Although Hubbard (1896) implied that he col-
lected this species at DeFuniak Springs, he was also
reporting on collections from Clearwater and may not
have actually had specimens from the former locality.
At least no such specimens have been discovered in his
collection, and all other records are from peninsular
localities. Blatchley (1928:10) recorded it from Lake
Worth, Enterprise, Sanford, Lake Mary, and Clearwater.
I have also seen specimens from Gainesville and Lake
Letta Subdivision (Highlands Co.).
BIOLOGY: This is one of the several obligates associ-
ated with burrows of the gopher tortoise. A few speci-
mens have been taken at light, but other than this,
nothing has been published on its habits except the
original observations by Hubbard (1894:305):
Specimens were found in every gopher
hole examined, and were frequently abun-
dant. Eighty-four specimens were collected
in a single burrow. The female forms food-
balls of gopher dung, after the manner of
related species above ground. In each of
these she lays a single egg, and then buries

Fig. 137-138. Head of Copris spp.: 137) C. minutus (Drury),
138) C. inemarginatus Blatch.

Copris gopheri Hubbard
Copris gopheri Hubbard 1894:305, 307, 310-311, Fig.

DIAGNOSIS: Small (length 7.5-10 mm), shining,
black, lightly punctate. Punctures everywhere sparser
than in any other species. Elytral striae sometimes
with a few rounded punctures basally, but without
coarse crenulate edges as in the other Florida species.
Although the elytral intervals are minutely punctate,
these are so small and scattered as to be almost un-
noticeable. Head horn of male weakly developed, never
elongate or curved as in minutus. Ninth elytral stria

it 4 or 5 inches deep in the sand beneath
the floor of the gallery. The material in
these balls is finely fibrous and dark green
in color. The larva begins eating near the
surface of the ball and forms a cavity
considerably larger than its body by press-
ing outward the dung, thus disturbing the
sphere and rendering it more or less pear-
shaped. In this operation it is evidently
assisted by the peculiar hump on the back,
so remarkably characteristic of the larvae of
this genus. The larva does not consume the
whole of its food supply, but disintergrates
the greater part of the mass, converting it
into a friable, black earth which falls away
at a touch. It finally constructs an oval
cocoon within the ball, with rather thin and
brittle walls formed from this black earth,
cemented by saliva or some other secretion,
and in this completes its transformations.
In the burrows which contain egg-balls,
specimens of the imago are less common,
and there appears to be a continuous suc-
cession of broods throughout the year.
I have not taken it in the several tortoise burrows
excavated at Gainesville nor in malt and proprionic
acid bait traps set in the entrances to these burrows.
The immature stages are unknown.

SPECIMENS EXAMINED: 17 from 4 Florida locali-
ties as follows: (3) Alachua Co., Gainesville, VII-65,
E. Gourley, Gopherus burrow. The following Gaines-
ville records are from blacklight traps with dates and
collectors as shown: (1) 15-V-68, R. E. Woodruff;
(1) 24-X-68, R. E. Woodruff; (2) 27-V-70, R. E.
Woodruff; (2) 24-X-71, F. W. Mead; (2) 3-4-X-72,
H. V. Weems, Jr.; (1) Highlands Co., Lake Letta Sub-
division, 16-VIII-61, T. Morris, blacklight trap; (4)
Pinellas Co., Clearwater, 27-VI, Hubbard (OSU); (1)
Seminole Co., Lake Mary (OSU).

SELECTED REFERENCES: Blatchley, 1928:10; Cas-
tle and Laurent, 1896:303; Hamilton, 1896:286; Hub-
bard, 1896:301; Matthews, 1961:4, 27, 31-32, 34, 36,
40, 56-59, Fig. 11, 14, 28-31, 35; Schaeffer, 1906:255;
Young and Goff, 1939:60-61.

Copris howdeni Matthews and Halffter
Copris howdeni Matthews and Halffter 1959:200-202.

DIAGNOSIS: Differs from all other U. S. species in
the unarmed head of both sexes. Similar in size (length
13-15 mm) only to inemarginatus in Florida, but that
species does not have the central notch in the clypeus.
Head entirely, evenly, densely punctate as the pro-
notum. Pronotum unarmed, median longitudinal sulcus
faint, barely visible on middle of the disc. Elytral
striae crenulate, intervals completely flat, densely and
coarsely umbilico-punctate. Most similar to fricator but
separated by the coarsely and densely punctate, rugose,
and flat elytral intervals.

TAXONOMIC NOTES: It is closely related to fricator
(Fab.), a common species in the northeastern U. S.

According to Matthews (1961:117) this species and
fricator cartwrighti Robinson represent isolated southern
populations apparently derived from fricator. He stated
that, "Its separation from fricator as a full species is
based primarily on its geographical isolation and ap-
pears justified on the basis of its great ecological and
climatic differences between its habitat and that of
Known only from Oneco and 3 mi. S. W. of Lake
Marion, both in central Florida.

BIOLOGY: Nothing is known of the biology or be-
havior, the five known specimens being taken at light
in March. The species is either very rare or occurs in
a restricted niche, possibly associated with some verte-
brate. I spent three nights at the type locality during
the same time of year that the types were collected,
operating a blacklight trap without success. The locality
is about five miles inland from the coast, at the rear
of an ornamental nursery operated by Miss Paula Dill-
man. This area contains a pond, bordered by fairly
dense hammock. Other Scarabaeinae taken in the
blacklight trap included Copris minutus and Pseudo-
canthon perplexus. Several nearby cow pastures were
examined without finding this species. Its habits are
still a great enigma.

SPECIMENS EXAMINED: One, the holotype, in the
U. S. National Museum, with the following data: Oneco,
Fla., 25-III-54, G. E. Ball, at light.
SELECTED REFERENCES: Matthews, 1961:34-35, 41,
108-109, 116-117, Fig. 11, 19, Table 2.

l d*A

Copris inemarginatus Blatchley
(Fig. 138)

Copris anaglypticus Say, Schwarz 1878:449 (misidenti-
Copris inemarginatus Blatchley 1918:54-55.

DIAGNOSIS: Medium sized (length 11-15 mm), bulky,
convex, shining, the clypeus without a central notch.
Easily distinguished from all other U. S. species by
the latter character. There is rarely a slight situation
in the clypeal margin where this notch should be, but
it is never indented and always without teeth. Upper
surface of head entirely punctate, area in front of the
horn almost rugosely punctate, punctures often coales-
cing. Head horn short, conical, and blunt, not differing
between the sexes. Pronotum unarmed, densely punc-
tate throughout. Elytral striae crenulate, punctures ap-
pearing transverse and quadrate; intervals convex, ap-
pearing smooth but very finely punctate. Pygidium
completely margined.

TAXONOMIC NOTES: This species was first recorded
under the name anaglypticus Say (Schwarz, 1878:449)
which is now considered a synonym of fricator (Fab.).
Blatchley described this species after some delay be-
cause he received the opinion from the late E. A.
Schwarz and H. S. Barber that this was " . . . merely
a depauperate form, due to biological conditions of
which we are now ignorant, but which is not specifi-
cally distinct from C. anaglypticus." There can be no
doubt about the distinctness of this species, and long
series have proven the constancy of the inemarginate
clypeus and other features. It was placed in the
"fricator complex" of species by Matthews (1961:110),
but it is isolated from the rest of the group by the
lack of the clypeal notch or emargination.

It is apparently confined to peninsular Florida with
records from Jacksonville to Miami. The type locality
is Dunedin, and Blatchley (1928:10) added Gainesville,
LaGrange, Lakeland, Port Orange, and Enterprise. In
addition to these, Matthews (1961:110) added High
Springs, Miami, Lutz, Tampa, 4 mi. N. W. Dunnellon,
Orlando, Kissimmee, Interlachen, 8 mi. S. Interlachen,
Sanford, 13 mi. N. O'Brien, Stemper, and Weeki
Wachee. Young (1959:103) reported an elytral frag-
ment from a Pleistocene deposit at Vero Beach. I have
seen it from 20 localities (for complete data see
Appendix 13).

BIOLOGY: This is a common species in cow dung
in the spring, but usually only in scrub habitats. Young
(1959:106) indicated that, along with the walking
stick (Anisomorpha buprestoides Stol.), this is a "highly
characteristic dung beetle" of the scrub. He went on to
state that " . . . perhaps Copris inemarginatus, a sort of
living fossil, did once utilize the dung of Equus, Mam-
mut, and Mammuthus which are also found as fossils
with it at Vero." Part of the type series was taken
from a dead turtle, and I have taken'one specimen on

the dried tendons of a horse carcass. Although Blatch-
ley (1918:55) reported specimens taken "at porch light
in June and July," I have never encountered it at light
nor in over 1000 blacklight trap samples examined.
I have taken a few specimens in traps baited with malt
and proprionic acid, and yeast. Matthews (1961:110)
stated that it is " . . . most active during the winter
and spring from November to April." I suspect that
the records reflect the time of year during which most
collecting has been done; the hot summer months are
not as appealing to the northern collectors as are the
mild winters. I have seen specimens collected every
month except September, October, and December.
My field notes contain the following observations:
at Redwater Lake (Putnam Co.) on Jan. 6, I collected
10 specimens from burrows beneath cow dung. In nearly
every case there was only a single specimen per dung
cake, the only exception being one cake with three
beetles. The burrows were slanting about 45 degrees
away from the source and were about six inches long.
No brood balls were found. They were more frequent
than C. minutus which is also abundant during the
winter. At Charlie Creek (Hardee Co.) on Jan. 21,
I took 15 specimens in burrows which were 6 to 9
inches deep. This was a turkey oak scrub area, but
the pasture was entirely open. On Mar. 20, near
Huntington (Putnam Co.), Dr. J. E. Lloyd and I took
93 specimens under cow dung in an open pasture
near a turkey oak scrub. Most of the specimens were
in the dung or in the top three inches of sand. Many
dung piles, seemingly in the proper condition, contained
no beetles, but over 40 specimens were taken in a
single pile. The immature stages are unknown.

SPECIMENS EXAMINED: 160, including the holo-
type, from 20 Florida localities, representing 27 collec-
tion records (for complete data see Appendix 13).

SELECTED REFERENCES: Blatchley, 1919:31; 1928:
10; Dozier, 1918:332; Matthews, 1961:35, 40, 108-110,
Fig. 11, 19, 47, 56, Table 2; Young, 1959:103-106.

Copris minutus (Drury)
(Fig. 136-7)
Scarabaeus minutus Drury 1770:78-79; P1. 35, Fig. 6.
Scarabaeus silenus Fabricius 1775:21.
Scarabaeus ammon Fabricius 1781:24.
Scarabaeus lar Fabricius 1787:13.
Copris reflexus Panzer 1794:7.
Copris minutus (Drury), Horn 1870a:42-51.
DIAGNOSIS: Small (length 8-12.5 mm), shining, black,
the head horn often well developed in both sexes. The
only Florida species which has a long, narrow head
horn in major males. Easily distinguished from all
other American Copris by the uniformly punctate proe-
pimeron. In all other species the proepimeron is
divided longitudinally by a feeble carina which dif-
ferentiates an outside, densely punctate area, from an
inside, almost impunctate one.

TAXONOMIC NOTES: The synonymy cited above was
confirmed by Matthews (1961). There is considerable
variation in size and development of the head horn. In
plotting horn height against femoral length, Matthews
(1961:Fig. 2) found that most specimens from Mobile
and Montgomery, Alabama, and Clarksville, Florida
fell near the upper extreme of the curve. However, two
specimens from that area fell near the bottom of the
curve. Further study of long series of specimens from
throughout the range will be necessary before any
interpretation can be made of this variability.
It occurs throughout the eastern U. S. from New
Hampshire south to Florida, west to Texas and north
to Iowa. In Florida it was recorded by Matthews from
36 localities from Miami in the south to Calhoun
County in the north. I have seen specimens from most
of the state, representing 75 localities. There are very
few records for the east coast and none from the
Everglades or the Keys.
BIOLOGY: This is a very abundant species in cow
dung throughout the year in Florida. It is also attracted
to lights, especially in the winter. Despite its abun-
dance, very little has been published on its behavior
or biology. Ritcher (1945:10-11) described the larva
and added the following observations:
Adults of C. minutus construct a brood cham-
ber several inches deep in the soil beneath cattle
droppings. Here several balls are formed from
an unshaped mass of dung. Balls with eggs
have a slight pyriform shape, measuring 13 to
15 mm in length and 13 to 14 mm in width.
Adults are usually found in the brood chambers

with the balls even after the larvae within
have pupated.
I have taken a few specimens on a dead dog and
on rotting blue crabs, but it appears to be rarely
necrophagous. Very few specimens have been taken
in numerous bait traps using fermenting malt.

SPECIMENS EXAMINED. Over 1,300, of which 943
were from 75 Florida localities, representing 209 col-
lection records (for complete data see Appendix 14).

SELECTED REFERENCES: Blatchley, 1910:916; 1928:
10; Brown, 1928a:25; Davis, 1966:213; Dillon and
Dillon, 1961:512, P1. 49, Fig. 9; Dozier, 1918:332;
1920:365; Horn, 1870a:42-51; Matthews, 1961:17, 31,
36, 40, 59-63, Fig. 2, 11, 14, 36, 51, 65, Table 2;
Miller, 1954:Table 1, 2; Ritcher, 1945:10-11, Fig. 36,
41 (larva); Schaeffer, 1906:255.

Genus PHANAEUS MacLeay
(Fig. 1, 145-54)
Phanaeus MacLeay 1819:124.

TYPE SPECIES: Phanaeus vindex MacLeay 1819
(designation not known, but cited by Matthews 1966:

DIAGNOSIS: Large and bulky (length 13-20 mm,
width 7-12 mm), brightly colored with metallic green
or blue, often with red or golden reflections. Males
have the pronotum flattened somewhat and projecting at
the posterior angles, and the head has a long curved
horn. The front tarsi are absent in the males (Fig.
49) but usually present in females. The elytral striae
feebly impressed, the intervals reticulately punctate.
Easily distinguished from all other Florida Coprini
by the bright color and absence of tarsi in the males
(this character found only in one other genus, Delto-
chilum, of the Scarabaeini).

TAXONOMIC NOTES: The genus was revised by
d'Olsoufieff (1924), but several species have been de-
scribed since then. Robinson (1948e) revised the U. S.
species, but his treatment is unsatisfactory in many
cases. Three species have been reported from Florida,
but one of these is based on an allopatric subspecies
of P. difformis which is here synonymized. Most of
the subspecies created by Robinson do not appear valid
to me. The entire genus is in need of revision, as are
our few U. S. species. Edmonds (1972) published an
excellent morphological study of the "phanaeine"
Scarabaeinae which will provide a firm basis for future
taxonomic studies.

proximately 100 species known, of which nearly all are
Neotropical, with 9 found in the Nearctic and a single
species known from the West Indies (endemic to
Jamaica). Matthews (1966:38) stated that the genus is
of South American origin with extensive penetration of



Fig. 145. Lateral view of Phanaeus vindex MacL., male (4X).

Central and North America, which it must have invaded
early (in the Upper Cretaceous or Eocene, according
to Halffter, 1964).
BIOLOGY: Most of the North American species are
coprophagous, but a high percentage of the South
American ones are necrophagous. A few species are
diurnal, but most are strictly nocturnal. Some
species are confined to forested areas, but several are
found in the grassland biome, especially those which
utilize herbivore dung. The behavior has not been
studied in detail for the Florida species.
Stewart (1967) studied the food preferences of
dung beetles in Georgia by using baited pitfall traps.
The most abundant species was Phanaeus vindex MacL.
with P. igneus a distant second. In three separate lo-
cations, feces preferences by the beetles were in the
following order: swine, opossum, dog, cow, raccoon,
and horse. None was found in chicken or lamb feces.
Cow dung was far less attractive than that of swine
or opossum " . even in an environment dominated
by cattle."
The larva is known for only one of the U. S.
species (vindex). It can be distinguished from the other
known Florida larvae of the subfamily Scarabaeinae
by the following combination of characters (Ritcher,
1945:12-13): prothoracic shield with an anteriorly
projecting, angular process on each side; glossa with a
transverse row of closely spaced spine-like setae an-
terior to the hypopharyngeal oncyli; venter of last
abdominal segment with a single broad, caudal, median
lobe; tarsal claws absent; medium portion of last ventral
abdominal segment covered with a large quadrate patch
of stout, caudally directed, spine-like setae; maximum
width of head capsule of third instar 4.2-4.6 mm.
SELECTED REFERENCES: Blanchard, 1885:167-169;
Blatchley, 1910:917; 1928:11; d'Olsoufieff, 1924:5-172;
Edmonds, 1972; Gillet, 1911:81-87; Matthews, 1966:

3845; Ritcher, 1945:12-13 (larva); Robinson, 1948e:
Key to the Florida species and subspecies
of Phanaeus
1. Elytral intervals one and two (and the sutural
one) carinate for at least one-half their length;
the remaining intervals with carinate reticulations
between the punctures which are elongate de-
pressions and never round (Fig. 146); major male
with head horn conical for its entire length (Fig.
145), not flattened or spatulate near the tip; pos-
terior pronotal angles produced . . . vindex MacL.
1'. Elytral intervals feebly convex, but never carinate,
the punctures usually rounded, the areas between
simply convex but not carinate (Fig. 147, 149);
major male with head horn flattened dorsoven-
trally for part of its length, expanded laterally
and somewhat spatulate near the tip (Fig. 1);
posterior pronotal angles not produced ........
.................. . . igneus M acL.. . . .2

2(1'). Elytra shining to the unaided eye, the first three
intervals with very few punctures except near the
suture; all elytral punctures smaller, less dense,
and less alutaceous (Fig. 149-50); peninsular
Florida ............... igneus floridanus d'Ols.
2'. Elytra dull to the unaided eye, the first three
intervals usually with some punctures throughout;
all elytral punctures larger, denser, and very
noticeably alutaceous (Fig. 147-8); western pan-
handle of Florida ........ igneus igneus MacL.

Phanaeus igneus igneus MacLeay
(Fig. 147-8)
Phanaeus igneus MacLeay 1819:133.
Phanaeus tityus Dejean 1836:155 (nomen nudum).
Phanaeus scabripennis Sturm 1843:106 (nomen nu-
Phanaeus igneus nigrocyaneus Gemminger and Harold
1869:1018 (nomen nudum, attributed to MacLeay
in Litt.).
Phanaeus igneus igneus MacL., Robinson 1948e:304.
DIAGNOSIS: Easily distinguished from the other
Florida species (vindex) by the characters in the key.
It can usually be told at a glance from igneus floridanus
by the dull, alutaceous elytra (Fig. 147-8). They are
rarely as brightly colored with red and bronze reflec-
tions as the peninsular form.
TAXONOMIC NOTES: Both subspecies were originally
described as full species, but since they appear to
intergrade somewhere in north Florida, I have con-
sidered them subspecifically distinct. Actually I have
seen no specimens which could not be separated, but
critical material is lacking from the northern and
western parts of the state. I have seen no specimens of
floridanus from north of the Florida peninsula. Al-

.uwffw-z r '- - -- , -
Fig. 146. Stereoscan photo of left elytron of Phanaeus vindex
MacL.: (17X). Compare Fig. 147-150.

though not a part of a formal synonymy, Edmonds
(1972:832) listed floridanus as a synonym of igneus.
There is a blue-black color form which appears to
be found on the fringes of the range (e.g., coastal
North Carolina and Georgia). This phenomenon has
been noticed in other species of Phanaeus, as well as in
igneus floridanus. This color form was responsible for
the name nigrocyaneus.

Outside of Florida it has been recorded from Alabama,
Georgia, and North Carolina. It was reported from
Clemson, South Carolina in the three papers by Cart-
wright (1934b, 1939a, 1950), although Kirk (1969:34
and 1970:31) reported it from several other areas. It
apparently does not occur in the mountains. In Florida
it appears to be confined to the panhandle west of the
Suwannee River, the easternmost record being Madison

BIOLOGY: The habits of this subspecies have not
been investigated. It is fairly common under cow and
human dung, and large numbers have been taken in
traps baited with malt and proprionic acid. I have
seen Florida specimens collected in March through
June, and September and October. The immature
stages are unknown.

SPECIMENS EXAMINED: 172, of which 67 were from
7 Florida localities as follows: (40) Calhoun Co.,
Clarksville, 21-III-54, H. F. Howden, malt traps; (1)
Jefferson Co., Monticello, 24-VI-33, F. W. Walker
[UMMZ]; (1) same data except 7-X-34, G. B. Fair-
child; (1) Leon Co., Tall Timbers Res. Sta., 14-IX-70,
D. L. Harris, pitfall; (1) same data except 8-XI-71;
(1) Liberty Co., Rock Bluff P. O., 6-IV-29, T. H.
Hubbell [UMMZ]; (6) Liberty Co., Torreya State
Park, 30-IV-46, F. N. Young [UMMZ]; (6) same data
except 12-IV-60; H. V. Weems, Jr., malt traps; (5)
Madison Co., 2-V-46, F. N. Young [ITMMZ]; (3)

Fig. 147-148. Stereoscan photos of left elytron of Phanaeus
igneus igneus MacL.: 147) 20X, 148) 128X. Note extent of
alutaceous areas.

Madison Co., Greenville, 2-6-IX-32, L. K. Gloyd
[UMMZ]; (2) Walton Co., DeFuniak Springs, 11-IX-
29, T. H. Hubbell [UMMZ].
SELECTED REFERENCES: Blanchard, 1885:169 (in
part); Blatchley, 1928:11 (in part); Brimley, 1938:199;
Edmonds, 1972:832; Miller, 1954:378-382, Table 1-2;
1961:735, Table 1; Robinson, 1948e:304.

Phanaeus igneus floridanus d'Olsoufieff
(Fig. 1, 149-50)
Phanaeus floridanus d'Olsoufieff 1924:94.
Copris floridanus Dols., Leng and Mutchler 1927:38
Phanaeus igneus floridanus d'Ols., Robinson 1948e:304.

Fig. 149-150. Stereoscan photos of left elytron of Phanaeus
igneus floridanus d'Ols.: 149) 24X, 150) 128X. Note extent
of alutaceous areas.

DIAGNOSIS: Bright, shining, metallic green, often with
red and bronze reflections. Easily separated from
typical igneus by the shining elytral intervals, the first
three often nearly impunctate (Fig. 149-50). Every-
where the sculpture less pronounced and punctures of
the elytral intervals usually rounded and not noticeably
alutaceous. Head horn of major male shown in Fig. 1.
TAXONOMIC NOTES: (see also this section under
the nominate subspecies). There is a blue-black form
with no coppery reflections, of which I have seen very
few specimens. All of these were from coastal areas
(e.g., Marco, Miami, Pompano Beach, Vero Beach),
except a doubtful specimen from a student collection
labeled "Gainesville."

The type locality is "Est-Florida: Doubleday, Foster-
St. Johns Bluff." It is apparently confined to penin-
sular Florida, my southernmost record being Miami and
the northernmost the type locality. I have seen no
specimens from west of the Suwannee River.
BIOLOGY: It is presumably similar to that of the
nominate subspecies. I have taken specimens commonly
in cow dung and once on a dead dog, but by far the
most specimens were taken in traps baited with malt
and proprionic acid. H. R. Dodge collected 109 speci-
mens in fermenting yeast bait (made up on 9-X) traps
at Gainesville, on the following dates, with numbers in
parentheses: 10-13-X (9), 14-16-X (33), 17-X (21),
18-X (22), 19-X (24). The bait was much more
productive after 4 days and continued to be attractive
after 10 days. Specimens have been collected every
month of the year in Florida. The immature stages
are unknown. This species and vindex are a part of
the regular diet of the burrowing owl in Florida.
SPECIMENS EXAMINED: over 660 from more than
40 Florida localities, representing 135 collection records
(for complete data see Appendix 15).
SELECTED REFERENCES: Blatchley, 1928:11 (ig-
neus in part); 1932:17, 50, 163; Dozier, 1918:332;
1920:365; Edmonds, 1972:832; Hamilton, 1894:252.

Phanaeus vindex MacLeay
(Fig. 145-6)

Phanaeus carnifex Linnaeus 1758:346 (in part, not
Phanaeus vindex MacLeay 1819:133.
Phanaeus vindex cyanellus Robinson 1938:107. (NEW
Phanaeus difformis magnificens Robinson 1948e:302.

DIAGNOSIS: Easily distinguished from igneus and
subspecies by the characters in the key. Pronotum of
the major male with a flattened area projecting pos-
teriorly at the posterior angles. In a few specimens
the elytra are blue and the pronotum green (cyanellus),
a striking contrast to the normal bronze or reddish
pronotum and green elytra.

TAXONOMIC NOTES: The name carnifex, long used
in American literature for this species, is properly
applied to a Jamaican species. The nomenclatural
problems have been thoroughly discussed by Barber
(1928) and Matthews (1966). The name cyanellus
was proposed by Robinson (1938:107) as a variety for
the blue-green form mentioned above. Color changes
on the margin of the geographic range are found in
several species of Phanaeus (e.g., torrens, quadridens,
igneus, etc.) but I do not believe that such forms
represent subspecies. In most cases there appear to be
no differences except color. The synonymy of difformis
magnificens is proposed after examination of the types.

Although I have found no other Florida specimens to
match exactly, I believe they fall within the range of
variation. It is likely that difformis difformis LeConte
is only a western subspecies of vindex, although I am
not formally proposing such at this time because of a
lack of material from the western U. S. This is a
wide-ranging species which has a great amount of
variation, and additional study will be required to
interpret this variability in light of geography and
behavior. Edmonds (1972) lists a index group, con-
taining 32 of the 44 species of Phanaeus, and the
vindex complex composed of 4 species indexe, diffor-
mis, igneus, and triangularis).

It is widely distributed throughout the eastern U. S.
from Massachusetts to Florida, west to Texas, and north
to South Dakota. In Florida it is found throughout the
state except for the Everglades and Keys.

BIOLOGY: Although this is a common, morphologi-
cally well known, widely distributed species, there have
been few detailed observations on its biology and
behavior. It is primarily coprophagous, with a decided
preference for human dung. The dung is provisioned
for the larvae in nearly vertical burrows a few inches
beneath the source. The dung wads are irregularly
shaped, but often pyriform, the egg being deposited in
the teat-like end. The adults, according to Ritcher
(1945:13), " . . . do not stay with the young as do
the species of Copris." For a characterization of the
larva see this section under the genus.
As is true with many coprophagous scarabs, a
phoretic association exists with several mites. This

association was studied by Stewart and Davis (1967),
who stated that Macrocheles amygdaligera (Berl.) and
" . . . probably all these phoretic macrochelids are
predators on insect eggs, small insect larvae, and
nematodes." They noted that in the laboratory these
mites readily fed on eggs of the swine stomach worm,
Physocephalus sexalatus (Molin), for which Phaneus
vindex is a known intermediate host. The beetles are
also known as intermediate hosts for a second stomach
worm, Ascarops strongylina (Rudolphi) (Stewart &
Kent, 1963). The close association or "consortisms"
between swine, nematodes, dung beetles, and mites
appears to be very complex. Future work in this regard
should prove very interesting. Specimens have been
taken every month of the year in Florida, but it does
not appear as common here as in Georgia. It is one
of the regular elements in the diet of the burrowing
owl in Florida, the remains of hundreds of beetles
being found in their pellets.
SPECIMENS EXAMINED: Over 215, of which 110
were from 45 Florida localities, representing 91 collec-
tion records (for complete data see Appendix 16).
SELECTED REFERENCES: Barber, 1928:383; Becton,
1930:315-321, Fig. 1-10 (alimentary tract); Blanchard,
1885:168-169; Blatchley, 1910:917, Fig. 363; 1928:11;
Borror and Delong, 1964:313, Fig. 22-57c, d; Brown,
1928a:25; Comstock, 1940:517; Dillon and Dillon, 1961:
513, P1. 50, Fig. 1-2, color P1. D; Edmonds, 1972;
Hardenberg, 1907:563, P1. 31, Fig. 22, P1. 34, Fig. 3
mouthpartss); Lindquist, 1933:111; Miller, 1961:738-
739; Ritcher, 1945:12-13, Fig. 7, 20, 24, 31, 33, 40
(larva); Stewart and Davis, 1967:20-26; Stewart and
Kent, 1963:158-159.

(Fig. 155-176)

The tribe is represented in the Western Hemisphere
only by the type genus Onthophagus. It can be dis-
tinguished readily from all other Florida Scarabaeinae
by the characters in the key. Eight other genera are
currently recognized in the tribe: Mimonthophagus
(Ethiopian), Macropocopris (Australian), Caccobius
(Palearctic, Oriental, Ethiopian), Milichus (Ethiopian),
Cyobius (Oriental), Anoctus (Oriental), Caccophilus
(Ethiopian), and Phalops (Ethiopian, Oriental).

Genus ONTHOPHAGUS Latreille
(Fig. 155-176)
Onthophagus Latreille 1802:141.
TYPE SPECIES: Scarabaeus taurus Schreber (by
DIAGNOSIS: Medium sized (length 2-8.5 mm), alu-
taceous to shining, pubescent or glabrous, color black
to metallic green or bronze, a few species maculate
with red, orange, or yellow spots. General shape oval,
usually somewhat flattened above. Clypeus entire or
notched, the margin raised or not evident. Head and/
or pronotum often furnished with horns or protuber-
ances in the male. Scutellum not visible. All tarsi
present. Middle and posterior tibiae expanded at the
apex, the middle with two spurs, the posterior with
one. Elytra with seven striae, not deeply impressed,
often broken into lines and dashes, the intervals not
strongly convex.


Fig. 155. Onthophagus p. polyphemi Htbbard, line = 5mm.

Most similar in general appearance to Oniticellus
(Fig. 177), from which it can be distinguished by the
nine-segmented antennae and the lack of a scutellum.

TAXONOMIC NOTES: This is perhaps the largest
genus of beetles known, with 1,500+ described species.
The species of America north of Mexico were recently
revised by Howden and Cartwright (1963). Numerous
subgenera have been proposed, especially for Old
World species. Halffter and Matthews (1966:254)
listed 18 subgenera, but these have not been applied to
the North American forms. Most of the Florida species
are readily recognized by the characters of the key, but
a few are difficult without comparative material. Prob-
lems of subspeciation exist in three of our species:
striatulus, polyphemi, and orpheus. These are dis-
cussed in detail under each species.
Curtis (1825) designated nuchicornis L. as the type
species, but, as Latreille listed only one species when
he described the genus (taurus Schreber), Curtis' desig-
nation is invalid.

species in the world, 114 are known from the Western
Hemisphere, 37 from the U. S. and Canada, and 11
species and 2 subspecies from Florida. They are not
recorded south of 400 south latitude in South America
or north of 55� north latitude in North America. Baltha-
sar (1963) stated that the genus is Ethiopian in origin,
and according to Halffter (1964a) it probably entered
North America from Asia via the Bering land bridge
early in the Cenozoic since there is moderate radiation
in South America.
Five species are recorded from the Antilles (Mat-
thews, 1966:8), two of which have been introduced
there. One of these is African, and the other is
widespread in Mexico and Central America. Both have
been introduced only on the island of Martinique. None
of these occurs in Florida, and none of the Florida
species is found in the West Indies. Curiously, another
African species (depressus) has been introduced acci-
dentally into Florida and Georgia.

BIOLOGY: In such a large genus, as might be expected,
there is a wide variety of food habits, although most
species are coprophagous. The Florida species are
mostly found in cow dung, although a few apparently
are found only in other kinds of dung. O. polyphemi
is confined to the burrows of the gopher tortoise, and
orpheus appears to be primarily associated with pack
rats (Neotoma spp.), at least in Florida. One species
(striatulus) is found almost exclusively in fungi. The
habits of aciculatulus are completely unknown. It prob-
ably occupies some unusual niche not yet discovered.
0. tuberculifrons is the most abundant species in
Florida, and it has been recorded from the widest range
of foods, including several kinds of dung, fungi, car-
rion, and decaying vegetable matter. Among the species
not known in Florida there are some which are myrme-
cophilous, saprophagous, sarcophagous, and copro-
phagous (sometimes exclusively on dung of one mam-
mal). At least two species are found only on bat guano
in caves.

Most of the Florida species are more abundant in
sandy soils, although two (concinnus and subaeneus)
usually are found in hardwood forests with deep leaf
mold. The gopher tortoise occurs only in very sandy
areas, and the obligate 0. polyphemi coincides with
this habitat. These burrows, as micro-caverns, offer
a cooler, moister, darker habitat than the surrounding
Several of the Florida species are attracted to arti-
ficial baits, but only depressus (introduced from Africa
and subgenerically different) has been taken at light.
Baits include malt extract, fermenting yeast, proprionic
acid, Staley's sauce bait, and amyl acetate.
In most of the species which have been studied,
the rate of development is rapid and, in at least some
of the species, there are several generations per year
in Florida. Adults of tuberculifrons have been found
during the entire year at Gainesville and are active
during the coldest weather. Other species, such as
concinnus and aciculatulus, appear to have a narrow
period of adult seasonal activity.
The immature stages are known for only two (penn-
sylvanicus and hecate) of the 11 Florida species. The
larvae that are known are typical Scarabaeinae and
"hump-backed" in appearance. The "hump" is accen-
tuated by a dorsal, conical, state protuberance. The
known larvae are very similar, with minute differences
in the venter of the last abdominal segment.
The behavior of our species is almost completely
unknown. Stridulation is not known in the genus.
Howden and Cartwright (1963:7) recorded the follow-
ing generalizations about the biology:

Three instars are present, each stage lasting from
8 days to 2 weeks. Development from egg to
general adult is rapid, usually taking only 5 or
6 weeks. Emergence of the adults from the pupal
cells may be delayed in hot dry weather and
during the winter. The burrows, made at the
edge of or under cow dung, are often 'twisted,
sometimes branched and vary in depth from 1
to 9 inches, depending on the species. The oval
brood cell, approximately 11/2 times as long as
wide and filled with dung, is formed nearly
horizontally at the end of the burrow or branch.
After the egg is fastened on end to the side of
a small cavity formed in the upper end of the
dung, the cavity wall is sealed with the same
material. The burrow may then be partly refilled
with soil and the egg and subsequent larva left
without further attention.

SELECTED REFERENCES: Blatchley, 1910:917-921;
1928:11-13; Boucomont, 1932:293-332; Boucomont and
Gillet, 1927:103-263; Brown, 1926:99-101; 1927a:128-
133; Horn, 1875:137-141; Howden and Cartwright,
1963:1-133, 84 Fig.; Howden, Cartwright, and Halffter,
1956:1-16; Ritcher, 1945:13-15 (larvae); Schaeffer,
1914:290-300. *

Key to the Florida species of Onthophagus
(Modified from Howden and Cartwright, 1963)

1. Disc of pronotum tuberculate or simply punc-
tate; setae not flattened if present; head and/
or pronotum often with horns, protuberances,
or carinae in the male; never attracted to
light ................................... 2
1'. Disc of pronotum closely, setigerously, an-
nularly punctate, the ring-like punctures sepa-
rated by less than their diameters; each punc-
ture anteriorly with a distinct tubercle, centrally
with a short, recumbent, flattened, yellow
seta; in Florida known only from Highlands
Co.; head and pronotum without secondary sex-
ual characters; attracted to light ...........
................... . . depressus Har.

2(1). Disc of pronotum smooth or distinctly punctate,
tubercles lacking on disc or, if present, less than
one-half the diameter of nearest puncture in
basal area .............................. 3
2'. Disc of pronotum tuberculate; punctures, if
present, vague with diameters approximately
equal to that of the tubercles .. .......... 12

3(2). Color uniform, rarely with humeral umbone or
entire elytra lighter in general specimens .... 4
3'. Pronotum and elytra decidedly different in
color, elytra bi-colored or spotted ........ 13
4(3). Disc of pronotum virtually impunctate, shiny
black or brownish black; males without horns or
protuberances on head or pronotum, 5-7 mm
in length; in gopher tortoise burrows ........
.......... (polyphemi and subspecies) .. 5
4'. Disc of pronotum distinctly punctate, vestiture
variable; males with or without horns or pro-
tuberances on head or pronotum, size variable
. ,.. . . . . . . . . . . . . ..... . 6

5(4). Elytral intervals with one or two rows of seti-
gerous punctures (Fig. 155), punctures often
with a tubercle at anterior margin; peninsular
Florida .................. p. polyphemi Hub.
5. Elytral intervals 2 and 4 with only a few
scattered setae, nearly impunctate; west of Apa-
lachicola River ..........................
............ p. sparsisetosus Howd. & Cartwr.

6(4'). Shining black, brown, blue, green, or cupreous;
more than 4 mm in length; pronotum lacking
numerous smaller secondary punctures ...... 7
6'. Dull or feebly shining, alutaceous, brown or
black, or if shining only 3 to 4 mm in length;
some with small secondary punctures on pro-
notum ................................ 10

7(6). Pronotum without basal margin; clypeal em-
argination, if any, not dentate on each side;
male with at least small vertical head horns
basally; size and color variable .......... 8
7'. Pronotum narrowly margined basally; clypeus
usually bidentate, the teeth low, triangular, and
well separated; less than 5mm in length; shin-
ing blackish green or coppery; without tubercles
or horns behind eyes; male with short conical
pronotal protuberance .... subaeneus (Beauv.)

8(7). Second and third elytral intervals with three
rows of setigerous tubercles; male with long,
slender, vertical horn above each eye; male pro-
notal protuberance rounded; color blue-black
to bronze, never bright green; in fungi ......
........ (striatulus and subspecies) ...... 9
8'. Second and third elytral intervals with one or
two rows of setigerous punctures or tubercles;
male pronotum with bifurcate protuberance
projecting above head; color bright green; pack-
rat droppings .......... o. orpheus (Panz.)

9(8). Elytral intervals alutaceous between tubercles
...... ........... . s. striatulus (Beauv.)
9'. Elytral intervals smooth and shining between
tubercles ............. .s. floridanus Blatch.

10(6'). Anterior edge of pronotal punctures with a
small shining tubercle; clypeus weakly biden-
tate; west coast, vicinity of Dunedin .........
................. . . aciculatulus Blatch.
10'. Anterior edge of pronotal punctures lacking tu-
bercles; clypeus truncate to rounded, never bi-
dentate; widely distributed .............. 11

11(10). Dull brownish to black with pronotal punctures
generally the same size, usually all with setae;
larger (length 3-5 mm)... .pennsylvanicus Har.
11'. Shining black, pronotal punctures of two sizes,
very small punctures lacking setae scattered
among the large punctures; smaller (length
2-4 mm) .............. oklahomensis Brown

12(2'). Pronotum bright shiny green or bluish; elytra
usually bicolored, green with yellow base and
apex ........... .......... concinnus Lap.
12'. Color uniformly dull black to grey, usually
with red-brown spots at apex of elytra .......
................. . hecate blatchleyi Brown

13(3'). Pronotum shining between punctures, not dull
brownish black; pygidium usually distinctly
punctate with at least apical half shining; male
with vertical horns on head ................
............. . ... . s. striatulus (Beauv.)
13'. Pronotum dull brownish black, alutaceous be-
tween setigerous punctures; pygidium shallowly
punctate, usually alutaceous at least to apical
third; male without horns on,head ...........
............. ...... . tuberculifrons Har.

Onthophagus aciculatulus Blatchley
Onthophagus alutaceus Blatchley 1919:31 (not Wiede-
mann 1823:14).
Onthophagus aciculatulus Blatchley 1928b:128.
Onthophagus aciculatus Blatch., Leng and Mutchler
1933:38 (misspelling).

DIAGNOSIS: Small (length 3.8-4.5mm), black, shining,
minutely alutaceous. Clypeus bidentate, the teeth weak-
ly developed, barely reflexed. Head with two tubercles
basally and single tubercle near the middle; these tu-
bercles conical, not highly raised. Pronotum weakly
produced antero-medially into a convex projection, not
actually in the form of a tubercle; punctures shallow,
margined with a minute tubercle anteriorly; marginal
line fine but complete. Elytra more noticeably aluta-
ceous than head and pronotum; striae fine, the punc-
tures shallow and inconspicuous; intervals flat, seti-
gerously punctate-tuberculate, the setae fine, yellow,
and arranged in one or two rows per interval. The
female is unknown.
It is similar in size and general appearance to
oklahomensis, pennsylvanicus, and subaeneus. It is
easily separated from the first two by the bidentate
clypeus, convex antero-median protuberance (male),
the shallow pronotal punctures, and the three conical
tubercles of the head. From the latter it can be dis-
tinguished by the more alutaceous surface, the less con-
spicuous pubescence, less prominent pronotal protuber-
ance (male), and three, rather than two, conical head

TAXONOMIC NOTES: Blatchley renamed his aluta-
ceus which was preoccupied by alutaceus (Wiedemann,
1823). The second and third supplement to the Leng
catalogue misspelled his new name asaciculatus. It is
known only from three male specimens.

It is endemic to Florida, where it is known from Dune-
din and Pasco County, although Kirk (1970) recorded
it erroneously from South Carolina.

BIOLOGY: Practically nothing is known about this
rare species. The holotype was taken "on the wing,"
Jan. 7. Howden and Cartwright (1963: 80) recorded a
second specimen collected at Dunedin by Blatchley on
Jan. 22, 1921. However, Blatchley (1928:13), a refer-
ence not cited by Howden and Cartwright, recorded the
second specimen as ". . . beaten from oak, Jan. 31."
The third recorded specimen was collected in Pasco
County without precise locality or habitat data on
March 20, 1957. It is likely that it is a winter or
early spring species which is found on the dung of
some obscure mammal. I have searched for it in the
vicinity of the type locality on three occasions without
SPECIMENS EXAMINED: Two (of the three known
specimens), including the holotype in the Blatchley
collection at Purdue University. The other specimen is
labeled Pasco Co., Fla., 20-III-57, H. V. Weems, Jr.

SELECTED REFERENCES: Blatchley, 1928:12-13;
Howden and Cartwright, 1963: 79-80, Fig. 8, 83-84;
Kirk, 1970.

Onthophagus concinnus Laporte
Onthophagus concinnus Dejean 1836:157 (nomen nu-
Ontophagus concinnus Laporte de Castelnau 1840:87
(misspelling of the genus for all 39 species, but
the page headings are correct).
Onthophagus viridicollis Sturm 1843:108 (nomen nu-
Onthophagus protensus Melsheimer 1845:134.
Onthophagus subaeneus (Beauv.), Horn 1875:130 (mis-
DIAGNOSIS: Large for the genus (length 5.2-8.1 mm),
bicolored green and yellow, the major male with an-
teriorly directed pronotal protuberance. Easily dis-
tinguished from all other Florida species by the yellow
and green color pattern. In sculpture it shares with
hecate blatchleyi the elongate pronotal tubercles or
TAXONOMIC NOTES: In most of the U. S. literature
this species is reported under the name subaeneus
(Beauv.). Howden and Cartwright (1963:112) pointed
out that this was a result of an early misidentification
by Horn (1875), and the name subaeneus is now ap-
plied to the species previously known in nearly all the
U. S. literature as cribricollis Horn.
Howden and Cartwright (1963:111) recorded it from
the following states: Florida, Georgia, Mississippi, New
Jersey, North Carolina, Pennsylvania, South Carolina,
and Tennessee. They listed the following Florida rec-
ords: Miami, Monticello, Mossyhead, Newmans (sic)
Lake, and Wacissa. Blatchley (1928) did not record it
from Florida. I have seen it from 4 additional Florida
localities (see specimens examined).
BIOLOGY: Howden and Cartwright (1963:111-112)
reported it as follows:
Biologically, O. concinnus is still some-
what of an enigma. It is a widely distributed
species, occurring in the spring and fall from
New Jersey to Florida and westward to
Mississippi. Nowhere does it seem common.
A possible explanation of its seeming rarity
may stem from the fact that the adults are
usually taken on the droppings of small
mammals, skunks, foxes, and possibly others,
only occasionally being taken on human
feces or cow dung. It has been taken under
fungi, under chicken manure, and a few spe-
cimens have been collected in fermenting
malt traps. Several live females were placed
in flower pots and supplied with fresh cow
droppings, but none of the specimens
evinced any interest in the dung, dying with-
out attempting to construct brood cells.

Several of the specimens that I have seen were
taken in human dung in forested (hammock) areas.
Others were taken in a Malaise-type trap in a for-
ested area, and several were taken in unbaited pitfall
traps at Tall Timbers Res. Sta. (Leon Co.). It appears
to be found in the same habitats as Glaphyrocanthon
viridis. Florida specimens have been taken from
March through September. The immature stages are

Fig. 156. Stereoscan photo of male pronotal horn of Ontho-
phagus hecate blatchleyi Brown (dorsal view, 56X).

SPECIMENS EXAMINED: 57 from Alabama, New
Jersey, and Georgia, as well as 56 from 4 Florida lo-
calities as follows: (1) Alachua Co., Gainesville, 12-
V-24, H. E. Bratley (determined as janus by Schwarz);
(1) Gainesville, 12-V-66, R. E. Woodruff, human
dung; (7) Gainesville, 10-IV-67, F. J. Moore, human
dung; (6) Gainesville, 17-XI-7-XII-72, H. R. Dodge,
feces bait traps; (1) Dade Co., Brickell Hammock, 7-
VII-36, F. N. Young; (37) Leon Co., Tall Timbers
Res. Sta., Woodyard Hammock, various dates from
27-III to 28-IX, D. L. Harris, unbaited pitfalls; (1)
Liberty Co., Torreya St. Pk., 1-IV-64, H. V. Weems,
Jr., Malaise trap; (1) same data except 5-VII-65; (1)
same data except 18-V-66, trapped in window.
SELECTED REFERENCES: Boucomont, 1932:329;
Brimley, 1938:200; Howden and Cartwright, 1963:108-
112; Lacordaire, 1856:109, Fig. 7, 61-63; Schaeffer,

Onthophagus depressus Harold

Onthophagus depressus Harold 1871a:116.
Onthophagus carter Blackburn 1904:147.

DIAGNOSIS: Large (length 6.0-7.7mm), oval, brown-
ish black to gray, dull. Clypeus bidentate, narrowly
emarginate between, the surface with transversely elon-
gate rugae or tubercles. Pronotal punctures variable in
size, but usually round, with an anterior tubercle, some-
times obscuring one-half of each puncture. Differs from
all other Florida species by the short, flat, broad setae
in each puncture of the dorsal surface (head, prono-
tum, elytra, and pygidium). Practically no secondary
sexual dimorphism, and no horns or protuberances
on head or pronotum in either sex.
TAXONOMIC NOTES: The above synonomy was sug-
gested by Arrow (In Cartwright, 1938:114). However,
no mention is made of the name carter or reference
to the synonymy in the recent revision by Howden
and Cartwright (1963). It is related to bituberculatus,
another African species, which was reported from
Martinique in the West Indies (Matthews, 1966:25).
Both species belong to d'Orbigny's (1913) 32nd group
of African species.
The type locality of depressus is Caffraria, southeast
Africa; that of carter is Australia. In the U. S. it has
been reported from Lyons, Vidalia, and Wenona,
Georgia and Lake Placid and Sebring, Florida (How-
den and Cartwright, 1963:127). It was first reported
from Florida by Robinson (1948c:177) based on a
specimen from Archbold Biological Station. Frost
(1963:34) erroneously considered his specimens the
first Florida record. I have seen additional specimens
from Lake Letta Subdivision, north of Avon Park, and
from Archbold Biological Station (both in Highlands
Co.). It is peculiar that an African species should turn

100A 0&r

up in such disjunct localities as Australia, Georgia,
and Florida. This is especially true since the U. S. lo-
calities are inland, and it has not been found near the
coast. It is doubtful that it occurs in the intervening
areas between the Georgia and Florida localities, since
it is attracted to light, and should be easily detected
with blacklight traps.
Matthews (1972:305-306) indicated that O.
depressus was accidentally introduced into Australia
(probably near Sydney) before 1900. Shortly after-
wards (Blackburn, 1904) it was described as 0. carter.
It appears that it did not move from the Sydney area
until after 1941 and may now be expanding its range.
Matthews (loc. cit.) mentioned that several specimens
were found in marine littoral conditions in Australia
which is ". . . undoubtedly an essential part of its
dispersal ability."
BIOLOGY: It is the only Florida species of Onthopha-
gus that is attracted to light. Although it was taken in
cow dung by P. W. Fattig in Georgia, I have searched
for it in vain in cow dung in Florida. At the Archbold
Biological Station, where it has been taken at light, I
examined several hundred piles of cow dung without
finding a single beetle. Specimens have been taken
from March through October in Florida. The immature
stages are unknown.
SPECIMENS EXAMINED: Over 1000, of which 140
were from Highlands Co., Florida as follows: (3) Lake
Placid, Citrus Tower, 3-VII-60, R. E. Woodruff, at
light; the remainder were from Lake Letta Subdivision,
by Ted Morris in blacklight trap, with numbers and
dates as follows: (1) 27-III-61; (37) 31-III-61; (9)

2-V-61; (2) 31-V-61; (13) 14-VI-61; (15) 21-
VI-61; (5) 27-VI-61; (3) 19-VII-61; (5) 8-VIII-61;
(40) 22-VIII-61; (2) 29-VIII-61; (1) 6-IX-61; (1)
2-X-61; (1) 19-IV-62; (2) 28-V-62.
SELECTED REFERENCES: Cartwright, 1938:114;
Frost, 1963:34; 1964:142; Howden, 1966a:1186; Fig. 18;
Howden and Cartwright, 1963:126-127, Fig. 4, 79-80;
Matthews, 1972:305-306; Paulian and Lebis, 1960:23,
Fig. 9; Robinson, 1948c:177.

Onthophagus hecate blatchleyi Brown
(Fig. 156)
Onthophagus hecate (Panz.), Blatchley 1928:12.
Onthophagus blatchleyi Brown 1929a:86-87.
Onthophagus hecate blatchleyi Brown, Howden & Cart-
wright 1963:120-123, Fig. 11, 73-75.

DIAGNOSIS: Large for the genus (length 5.5-8.2 mm),
dull, gray, usually with a few orange to red spots at
the elytral apices. In the elongate pronotal granules or
tubercles it is similar only to concinnus in Florida.
However, that species is bicolored green and yellow.

TAXONOMIC NOTES: Howden and Cartwright (1963:
122), in relegating this form to subspecific status, re-
marked that it ". . . has been treated at a subspecies
not because it is lacking in distinguishing character-
istics, but because all the characteristics in specimens
from north of peninsular Florida appear to blend with
the true hecate." I have not seen any Florida specimens
which are intermediate or similar to the nominate
subspecies. However, critical material is lacking from
the western panhandle and from the zone of reported
intergradation. The Florida population nearly always
has the orange spots at the elytral apices, the pronotal
granules are less elongate and more widely spaced, and
the dorsal pubescence is shorter and less conspicuous.
The specimens I have seen of hecate from parts of the
range (e.g. Missouri and Kansas) differ in some of
the same relative characters (e.g. denseness of pro-
notal granules) from the northeastern populations.
This common, widespread species would make an ideal
subject for a detailed study of variation and subspecia-

The typical form is known only from peninsular Flori-
da. Howden and Cartwright (1963:Fig. 119) showed
intergrades from Clarksville, Florida through south-
eastern Georgia to the coastal plains of South Carolina.
Kirk's record (1969-70) from South Carolina probably
refers to these intergrades.
In Florida I have seen it from nearly all areas of
the peninsula, excluding the Keys. Although this sub-
species (or possibly intergrades with hecate) probably
occurs throughout the panhandle, I have seen only a
single specimen from that area (Jackson*Co.).

BIOLOGY: Howden and Cartwright (1963:123) re-
corded the following information: ". . . at cow dung,
small animal droppings, decaying fruits, and the fer-
menting malt-proprionic acid mixture. Specimens were
taken both in the sandhill areas and in the low ham-
mock areas of south Florida." In addition I have seen
specimens from dead fish, rotting pork, rabbit pellets,
rotting palm, human dung, deer droppings, dead dog,
and Steiner trap. Unbaited pitfall traps at Tall Timbers
Res. Sta. (Leon Co.) have produced 681 specimens. It
is not attracted to light. I have seen Florida specimens
collected every month of the year. The immature stages
are unknown.
SPECIMENS EXAMINED: 1,160 from 54 Florida lo-
calities, representing 362 collection records (for com-
plete data see Appendix 17).
SELECTED REFERENCES: Except for Kirk's (1969-
70) questionable South Carolina record, there are no
references other than those cited under the synonymy

Onthophagus oklahomensis Brown
Onthophagus oklahomensis Brown 1927a:128.
DIAGNOSIS: Small (length 2-4.1 mm), black, mostly
shining, somewhat flattened dorsally. Clypeus rounded
or truncate, never notched or dentate. Carinae of the
head weakly developed or obsolete. Head and prono-
tum without horns or protuberances in either sex. Most

similar to pennsylvanicus, it is distinguished by the two
sizes of pronotal punctures (interspersed), generally
smaller size, shinier pronotum, and more pronounced
posterior pronotal margin.
TAXONOMIC NOTES: Although very close to pennsyl-
vanicus, it is distinct in detailed morphology as well
as habitat preferences. It is the smallest U. S. species
(length 2 mm).
It was recorded by Howden and Cartwright (1963:81)
from the following states: Arkansas, Florida, Georgia,
Kansas, Louisiana, Mississippi, North Carolina, Okla-
homa, South Carolina, Tennessee, Texas, and Vir-
ginia. In Florida they listed it as statewide (25 locali-
ties). I have seen specimens from most of Florida ex-
cept the Everglades and the Keys.
BIOLOGY: Howden and Cartwright (1963:82) reported
the following:
The habitat preference of 0. oklahomen-
sis differs from pennsylvanicus, for though
both species are sometimes taken together,
the former is restricted almost entirely to
sandy areas. It is commonly taken in the
sandhill regions of the southeastern coastal
plain as well as in sandy areas in Okla-
homa and Texas. The species appears to be
generally a dung feeder, making shallow
1-to-3-inch burrows under or beside piles
of cow dung. Besides being attracted to
dung, adults come readily to rotten melon
rind, bananas, and malt and proprionic acid
traps. In the laboratory, cow manure was
used by the beetle for construction of small

oval cells buried 1-to-2 inches deep in
packed sandy clay. Sand grains coating the
cells made them difficult to measure, but 11
cells averaged approximately 10 mm. long
by 8 mm. wide. In this species as in pennsyl-
vanicus, development from egg to adult
takes about 3 weeks or slightly longer. Sev-
eral of the cells were formed about June
25, and on July 17 some contained pupae or
general adults. The length of the various
instars was not ascertained.
I have seen specimens with the following habitat
information: human dung, cow dung, dog dung, fleshy
fungus, Jap beetle trap, and bait traps with malt, pro-
prionic acid, asafoetida, and yeast. It is not attracted
to light. Specimens have been taken in Florida every
month except January, February, and September. Al-
though several hundred hecate blatchleyi were taken
in unbaited pitfall traps at Tall Timbers Res. Sta.
(Leon Co.), only 1 oklahomensis was taken in the same
SPECIMENS EXAMINED: Over 600, of which 540
were from 23 Florida localities, representing 60 collec-
tion records (for complete data see Appendix 18).
SELECTED REFERENCES: Boucomont, 1932:319;
Howden and Cartwright, 1963:80-82, Fig. 3, 58; Knaus,

Onthophagus orpheus orpheus (Panzer)

Scarabaeus orpheus Panzer 1794:5.
Onthophagus orpheus (Panz.), Sturm 1843:107.
Onthophagus janus var. orpheus (Panz.), Horn 1875:
Onthophagus orpheus orpheus (Panz.), Howden and
Cartwright 1963:47-50.

DIAGNOSIS: Large for the genus (length 5-9 mm),
shining green or bronze. Male with a deeply forked,
anteriorly projecting protuberance on the pronotum and
two dorsally projecting, short horns on the base of the
head. Clypeus transversely rugose and punctate. Pro-
notum punctate-tuberculate, the tubercles projecting
posteriorly at the anterior edge of each puncture;
punctures bearing fine, whitish setae, more noticeable
at the sides. Distinguished from the other two sub-
species of orpheus by the pronotal and elytral punctures
with tubercles on their anterior margins. It can be
separated from the other Florida species by the shape
of the male pronotal protuberance and the other char-
acters in the key.

TAXONOMIC NOTES: Apparently all Florida speci-
mens are referrable to the nominate subspecies. This
polymorphic species is in need of further study. Howden
and Cartwright (1963:49) stated that ". . . instead of
being divisible into several subspecies as treated here,
[it] may represent a series of sibling species; how-
ever, the taxonomy of the group will remain obscure

.016 0

until a great deal more is known about the biology of
the complex." Most of my Florida specimens were
confirmed as this subspecies by Cartwright.

It was recorded by Howden and Cartwright (1963)
from the following states: Florida, Illinois, Indiana,
Iowa, Kansas, Maryland, Minnesota, New Jersey, New
York, Ohio, Oklahoma, Pennsylvania, South Carolina,
Tennessee, Texas, Virginia, and Wisconsin. They listed
the following Florida localities: Enterprise, Lake City,
Live Oak, Florida Caverns State Park, and Key Largo.
Blatchley (1928:12) also listed Crescent City.

BIOLOGY: All of my Florida specimens have been
associated with woodrats (Neotoma floridanus and sub-
species). I have taken it also in Ohio in woodrat
droppings. Howden and Cartwright (1963:50) recorded
the following hosts: buzzard's nest, woodchuck burrows,
in a cave, cow dung, and malt traps. It is not attracted
to light. They suggested that, "Inasmuch as most of the
species of Onthophagus that are not general dung feed-
ers seems to have a very restricted host preference, it
seems quite likely that further investigation may show
that three or four morphologically similar species
with quite diversified habits are placed here under the
name orpheus."

SPECIMENS EXAMINED: 40, of which 21 were from
2 Florida localities as follows: (19) Jackson Co., Flor-
ida Caverns St. Pk., 6-X-60, R. E. Woodruff, Neotoma
droppings; (1) same data except 18-IV-63; (2) Monroe
Co., Key Largo 7-VI-60, R. E. Woodruff & L. J.
Bottimer, in dung chamber in nest of Neotoma floridana


small; (1) same data except 7-XII-66, R. E. Woodruff,
B. K. Dozier, & J. H. Knowles.
SELECTED REFERENCES: Blatchley, 1910:910; 1928:
12; Boucomont, 1932:311; Brown, 1926:100; Dillon and
Dillon, 1961:514-515, PI. 49, Fig. 6; Howden and Cart-
wright, 1963:47-50, Fig. 6, 26-27; Schaeffer, 1914:

Onthophagus pennsylvanicus Harold

Onthophagus ovatus (Linn.), Melsheimer 1806:4 (mis-
Onthophagus moeris Sturm 1826:178 (nomen nudum).
Onthophagus pennsylvanicus Dejean 1836:158 (nomen
Onthophagus pennsylvanicus Harold 1871a:115.
Onthophagus falcipes Harold 1871a:115.
Onthophagus pensylvanicus Har., Horn 1875:141 (mis-
Onthophilus pensylvanicus Har., Hubbard and Schwarz
1878:655 (misspelling of both genus and species).

DIAGNOSIS: Small (length 3.3-5 mm), black to dark
gray, dull to feebly shining. ClypeuS truncate to
rounded, not dentate. Pronotum without horns or
protuberances; punctures shallow, and nearly always
setigerous, separated by about one diameter, mostly of
same size, rarely with a few finer ones scattered near
the midline; the surface between the punctures finely
alutaceous. Elytral intervals dull, alutaceous, at least
third, fourth, and fifth with tubercles arranged in two
irregular rows. Most similar to oklahomensis, from
which it differs by the pronotal punctures being nearly
uniform with only rarely smaller ones interspersed. In
addition it is generally larger, less shining, and the pos-
terior pronotal margin is poorly developed.

TAXONOMIC NOTES: Although superficially similar
to oklahomensis, it is distinct both in morphology and
habits. The reference above to ovatus is presumed to be
a misidentification for this European species. Howden
and Cartwright (1963:84) indicated that ovatus is not
known from North America. Several early records for
pennsylvanicus could refer also to oklahomensis.
Howden and Cartwright (1963:84) recorded it from
"Colorado, South Dakota, and all states east and south
of these except New Mexico, Vermont, and Maine." In
Florida Blatchley (1928:12) recorded it from Sanford,
Sarasota, Dunedin, and Palmdale. I have seen speci-
mens from most of the state except the southern one-
fourth of the peninsula. Although records are lacking
for the entire eastern coast, this is probably an artifact
of collecting.
BILOGY: It has been found on dung of a variety of
animals. It is also sometimes found in carrion, rotting
fungi, and decaying fruits. It is not attracted to light.
It is much less common in Florida than in states to

the north. Howden and Cartwright (1963:85) stated
that specimens are ". . . most commonly collected in
areas having a clay-type soil, but also occur in fairly
sandy localities." I collected 63 specimens in cow dung
on Payne's Prairie near Gainesville in pure sand along
with oklahomensis and hecate blatchleyi.
Howden and Cartwright (1963) made the following
observations on the life history: winding burrows, two
to three inches deep, are made beneath and at the
margin of a pile of cow dung, where they terminate in
a cell averaging 6 mm wide by 10 mm long; a single
elongate egg is laid upright in a small cavity in the
upper end; development from egg to adult is approxi-
mately three weeks.
The larva has been described by Ritcher (1945 and
1966), but it is inseparable from that of oklahomensis.
It differs from hecate by having the raster with less
than 60 short, stout setae, usually in one patch, and
the maxillary stridulatory teeth number four to seven.

SPECIMENS EXAMINED: Over 300, of which 191
were from 18 Florida localities, representing 27 collec-
tion records (for complete data see Appendix 19).
SELECTED REFERENCES: Blatchley 1910:920; 1928:
12; Brown, 1926:100; Dillon and Dillon, 1961:515, P1.
49, Fig. 12; Howden and and Cartwright, 1963:82-85,
Fig. 2, 59-60; Lindquist, 1933:111, 120; Miller, 1954:
379-380, Table 1-2; 1961:738; Mohr, 1943:296; Ritcher,
1945:15, Fig. 45 (larva); Schaeffer, 1914:297; Wilson,
1932:79, 81.

Onthophagus polyphemi polyphemi Hubbard
(Fig. 155)
Onthophagus polyphemi Hubbard 1894:311-312, Fig.
Onthophagus polyphemi polyphemi Hubbard, Howden
and Cartwright 1963:35-38.
DIAGNOSIS: Large (length 4.7-6.9 mm), dark reddish
brown to black, shining. Clypeus rarely, barely emar-
ginate, truncate anteriorly; posterior portion delimited
by a transverse, evenly elevated carina. Another trans-
verse carina at the vertex of the head, more highly
elevated laterally (both carinae weakly developed in
the female). Head punctures setigerous and scattered
unevenly. Pronotum of major males swollen and slightly
produced anteriorly, that of the female evenly convex;
disc shining, without coarse punctures and setae; coarse,
setigerous, tuberculate punctures confined to lateral
area, and in males, to the area in front of the swelling.
Elytral striae vaguely punctate, intervals smooth, shin-
ing, and, except for the sutural one, each with a double
row of minute tubercles having setigerous punctures at
their bases.
It can be separated easily from all other Florida
species by the shining surface, and the pronotum is
without coarse punctures and setae over the central
one-third. It is very similar to the subspecies spar-

sisetosus but can usually be distinguished by the
characters in the key. In addition, the surface through-
out has more punctures, setae, and tubercles. One
character which appears to consistently separate the
two is the nearly impunctate second elytral interval
in sparsisetosus.
TAXONOMIC NOTES: Howden and Cartwright (1963:
40) considered the two forms subspecifically distinct
because ". . . the dorsal punctures, slightly reduced in
size and number, of specimens north (Tillman, S. C.)
and west (High Springs, Fla.) of Crescent City (type
locality) indicate possible intergradation."
The nominate subspecies is recorded from southeastern
South Carolina (Tillman) to south Florida (Miami),
and, along with sparsisetosus, is probably coextensive
with the range of the gopher tortoise (Gopherus poly-
phemus Daudin). The type locality is Crescent City,
Florida, and it has been reported from the following ad-
ditional localities: 4 mi. N. of High Springs, Gainesville,
Leesburg, Lutz, Miami, Stemper (Howden and Cart-
wright 1963:37); Lake Worth, Enterprise, Sanford,
Lake Mary, LaGrange, Funiak, Clearwater (Blatchley,
1928:12). The "Funiak" (probably DeFuniak Springs)
locality probably refers to sparsisetosus. Although re-
corded from Miami by Howden and Cartwright (1963:
37), this record is not shown in their distribution map
(Fig. 5) or reflected in their general statement of
the southernmost Florida record as "central Florida."
The distribution of the gopher tortoise is shown in
Fig. 67-8, based on unpublished data supplied by
W. A. Auffenberg.

BIOLOGY: Both subspecies of polyphemi are part of
an assemblage of arthropods which are obligates in the
burrows of the gopher tortoise. They feed on the dung
of the tortoise and are sometimes abundant in a single
burrow. Hubbard (1894:305) stated: "I did not find
this beetle in the few galleries examined in winter,
and it was probably in pupa at that season. In July
it was not rare. One of the burrows produced twenty-
one specimens. Its larva was not seen." Howden and
Cartwright (1963:37) reported specimens collected
in March, June, July, and August, with those from
March appearing freshly emerged. I have seen speci-
mens from these months as well as February, May,
and June.
Although a few specimens have been taken in malt
and proprionic acid traps set in the burrows, excava-
tion appears to be the only way to collect numbers
of specimens. The tortoise burrows vary considerably
in length (up to 20 ft.) and maximum depth (8 to
12 ft.). Excavation of such a burrow, especially being
careful to collect the arthropods, is a time consuming
and laborious undertaking. Hubbard (1894:303) stated
that one of his excavations ". . . was in loose yellow
sand of our pine woods subsoil, and when my explora-
tion was completed, so large a pit had been dug that
a coach and span of horses might have been swallowed
up in it." There have been no observations on the
behavior of the species, and the immature stages re-
main unknown.
SPECIMENS EXAMINED: 30 from 10 Florida local-
ities as follows (all were taken from gopher tortoise
burrows): (1) Alachua Co., 2 mi. W. Newnan's Lake,
8-VIII-62, R. E. Woodruff & B. Benesh; (1) same data
except 11-V-63, J. F. Anderson; (1) Alachua Co.,
Gainesville Airport, VII-65, E. Gourley; (1) same data
except 14-V-66; (3) same data except 27-VII-66; (1)
Alachua Co., Archer, 28-III-60, R. E. Woodruff; (6)
Gilchrist Co., Trenton, 28-IV-66, E. Gourley; (2) Lake
Co., Leesburg, 2-VII-38, C. C. Goff; (8) Leon Co.,
Tallahassee, 24-II-68, R. E. Woodruff; (1) Marion Co.,
1-VII-60, B. Papy; (1) Putnam Co., 12-VI-60, H. V.
Weems, Jr.; (2) Putnam Co., Crescent City, VII, Hub-
bard (paratypes); (1) Putnam Co., 4 mi. S. of Inter-
lachen, 29-III-60, R. E. Woodruff.
SELECTED REFERENCES: Blatchley, 1928:12; Castle
and Laurent, 1896:303; Howden and Cartwright, 1963:
35-38, Fig. 5, 14-15; Howden, Cartwright, and Halffter,
1956:10; Hubbard, 1896:301; Schaeffer, 1914:293;
Young and Goff, 1939:61.

Onthophagus polyphemi sparsisetosus
Howden and Cartwright

Onthophagus polyphemi sparsisetosus Howden and
Cartwright 1963:38-41, Fig. 5, 16-17.
DIAGNOSIS: Very similar to p. polyphemi except that
dorsally it has fewer punctures, setae, and tubercles.

The one character which appears to consistently sep-
arate the two is the nearly impunctate second elytral
interval in sparsisetosus. There are very few coarse
pronotal punctures at the sides and in front of the
protuberance in the males; this difference is less notice-
able in the females.
TAXONOMIC NOTES: (see this section under the
nominate subspecies).
The type locality of this subspecies is 6 mi. S. W. of
Stapleton, Alabama; paratypes are from 6.5 mi. S. of
Lucedale, Mississippi, near Clarksville, Florida, and
Funiak (DeFuniak Springs?), Florida. Howden and
Cartwright (1963) postulated that the Apalachicola
River might be the barrier between the subspecies.
I have seen a single specimen from Torreya State
Park, on the east side of the Apalachicola, and it agrees
well with sparsisetosus. If there is a natural feature
which marks the geographic division of the two forms,
then it must be something other than the Apalachicola
River. Critical material is not available from the
northern parts of the gopher tortoise range.
BIOLOGY: Practically nothing is known of the habits
of this subspecies, Jexcept that it is found in burrows
of the gopher tortoise. Presumably its habits are simi-
lar to those of the nominate subspecies. The type series
of 103 specimens from Alabama was taken in a single
burrow from one to 12 feet inside the entrance.
SPECIMENS EXAMINED: 35, of which 34 were
Alabama paratypes, and one was from Florida as fol-
lows: Liberty Co., Torreya St. Pk., 24-IV-61, H. V.
Weems, Jr.

SELECTED REFERENCES: Nothing has been pub-
lished on this recently described subspecies except the
original description.

Onthophagus striatulus floridanus Blatchley
Onthophagus nigrescens Blatchley 1916:94 (not d'Or-
bigny 1902:21).
Onthophagus floridanus Blatchley 1928b:128.
Onthophagus striatulus floridanus Blatch., Howden and
Cartwright 1963:45-47.
DIAGNOSIS: Large (length 5.5-7.2 mm), black, shin-
ing, punctures anteriorly tuberculate and setigerous.
Major male with two vertical, long horns at the base
of the head which fit on each side of the pronotal
protuberance. No other Florida species has such head
horns, except weakly developed ones beneath the elon-
gate pronotal protuberance in orpheus. Distinguished
from nominate striatulus by the fairly uniform black
color, more shining dorsal surface, and elytral intervals
having alutaceous sculpture between the tubercles. In
most specimens seen there are four rather than three
rows of tuberculate, setigerous punctures on each
elytral interval. Elytra never bicolored as sometimes
in typical striatulus.
TAXONOMIC NOTES: I have referred all Florida
specimens to this subspecies and have seen no speci-
mens that I consider intermediate with striatulus.
Howden and Cartwright (1963:46) stated:

*C gJe

0. floridanus Blatchley is subsequently listed
and described as an allopatric subspecies. Many
of the characteristics of s. floridanus, such as
tuberculate-punctate pronotum and black color,
appear separately in populations of s. striatulus,
but of the many specimens examined none with
a range outside the southeastern coastal plain
exhibited all the characteristics of s. floridanus.
This fact coupled with the restricted distribution
of s. floridanus would seem to make valid its
recognition as a subspecies.

This complex of forms is in need of further study.
Florida specimens were sent to Cartwright for inclusion
in the revision of Onthophagus (Howden and Cart-
wright, 1963). In a series from Gainesville, taken in
a single fungus, specimens were determined both as
striatulus and floridanus. I have reexamined these
and can find no trace of the alutaceous elytral sculp-
ture and therefore have no hesitation in referring them
all to floridanus. Howden and Cartwright (1963:46)
mentioned other specimens from Dunedin and High
Springs with varying amounts of faint alutaceous sculp-
ture on the elytra, and they stated: "These should
perhaps be considered intermediate forms." If these
forms are allopatric as these authors have stated, I
find it difficult to conceive of "intermediate forms"
from Dunedin, the type locality of floridanus.

Howden and Cartwright (1963:Fig. 1) showed typical
floridanus confined to the Florida peninsula. No speci-
mens have been seen by these authors or by myself
from the western panhandle. Although, as mentioned
above, intermediate forms were mentioned from Dune-
din, Gainesville, and High Springs, these are not in-
cluded in the zone of intergradation shown on their
map (Fig. 1).

BIOLOGY: Florida specimens have been collected
every month except March, April, July, and December.
This is the only Florida species which is primarily
an inhabitant of fleshy "toadstool" fungi. There is one
record from a dead opossum, and, probably like
s. striatulus, it will rarely be found in dung and other
carrion. Specimens have been trapped in bait cans
using karo syrup, malt, yeast, and asafoetida. Several
were taken in unbaited pitfalls at Tall Timbers Res.
Sta. (Leon Co.). At Gainesville I placed several fresh
"toad stool" fungi in a can about half full of sand,
and between Aug. 9 and 16, I collected 70 specimens
as follows: after 3 days (12), after 4 days (5), after
5 days (14), and after 7 days (39). It is not attracted
to light. The immature stages are unknown.

SPECIMENS EXAMINED: 197 from 16 Florida local-
ities, representing 57 collection records (for complete
data see appendix 20).

SELECTED REFERENCES: Blatchley, 1928:12; How-
den and Cartwright, 1963: 45-47, Fig. 1, 24-25.

Onthophagus subaeneus (Beauvois)
Copris subaeneus Palisot de Beauvois 1811:105.
Onthophagus subaeneus (Beauv.), Haldeman and Le-
Conte 1853:54.
Onthophagus cribricollis Horn 1881:76.
DIAGNOSIS: Small (length 3.3-5mm), head and
pronotum shining, dark iridescent cupreous to green;
elytra dull shining, black with cupreous or green cast.
Clypeus abruptly reflexed and broadly emarginate an-
teriorly, angulate on each side of the emargination,
often appearing bidentate. Pronotum completely mar-
gined; with a small conical protuberance medially,
barely extending over the anterior pronotal margin in
the major male; punctures spaced mostly less than two
diameters apart, anteriorly and laterally with tubercles
on their anterior margin, and all setigerous with fine,
long, whitish setae. Elytral intervals minutely aluta-
ceous, usually each with two rows of tuberculate, seti-
gerous punctures. It can be distinguished from the
other Florida species by the characters in the key. In
addition, the pronotal pubescence is finer and longer
than our other species. In the bidentate clypeus it is
most similar to aciculatulus, but that species is black,
has the pronotum alutaceous, and has three low conical
protuberances on the head.
TAXONOMIC NOTES: The name subaeneus has been
misapplied in most North American literature to the
species treated here as concinnus, following Howden
and Cartwright (1963). The present species had gone
under the name cribricollis since 1881.
Howden and Cartwright, (1963:75) recorded it from

-^ *

the following states: Alabama, District of Columbia,
Florida, Georgia, Illinois, Indiana, Kansas, Maryland,
Missouri, New Jersey, North Carolina, Ohio, Oklahoma,
Pennsylvania, South Carolina, Tennessee, Texas, and
Virginia. They listed Levy Co. and Gainesville as the
only Florida records. It was not recorded from the
state by Blatchley, and I have seen only one Florida
specimen (see Specimens Examined).
BIOLOGY: This species is rare in collections, al-
though apparently sometimes common in certain situ-
ations. Sim (1930:141) collected 120 specimens during
two seasons at Rancocas Park, New Jersey on rabbit
pellets. These were nearly all taken in June and July
in the sandy "pine-barren" country. He made the
following behavioral observations:
The beetles were most active on warm sunny
days after showers, and practically all were
found between 9 A.M. and noon. None was ever
observed on the wing in the afternoon or on a
cloudy morning. As in all species of Ontho-
phagus whose habits are known to me, cribricol-
lis buries its food where found and sinks it
vertically to a depth of a few inches, where the
subsequent grub lives in a double walled plaster
cell of its own manufactures. The entire meta-
morphosis was found to require about one
month. As in other species, this beetle, probably
overwinters as a hibernating adult buried singly
at a depth of several inches.
Howden and Cartwright (1963:75) recorded speci-
mens from fungi, carrion, under dung of various ani-
mals (including chicken manure), and malt or malt
and proprionic acid traps. Brown (1926) found it in
moist woodlands in Oklahoma. The immature stages
are unknown.
SPECIMENS EXAMINED: Six, one of which was
from Florida: (1) Leon Co., Tall Timbers Research
Station, 30-V-6-VI-70, D. L. Harris, pitfall trap in
Woodyard Hammock.
SELECTED REFERENCES: (all of these are under
the name cribricollis except Howden and Cartwright).
Blatchley, 1910:920; Brown, 1926:100-101; 1928a:26;
Howden and Cartwright, 1963:72-76, Fig. 8, 54-55;
Schaeffer, 1914:297; Sim, 1930:140-141.

Onthophagus tuberculifrons Harold
Onthophagus tuberculifrons Sturm 1843: 108 (nomen
Onthophagus tuberculatus Gemminger and Harold
1869:1038 (nomen nudum, attributed to Zimmerman
in litt.).
Onthophagus tuberculifrons Harold 1871a:115.

DIAGNOSIS: Small (length 3-5.5 mm), dull, aluta-
ceous, brown to black, elytra maculate with variable

orange spots, at least some present at the humeri and
apices. Clypeus broadly emarginate (males) or acutely
emarginate, the angles each side dentate (females).
Head with two "bumps" or tubercles at the vertex;
clypeal carina short, low, and most noticeable in fe-
males. Pronotum margined anteriorly and laterally,
but feebly so posteriorly; unmodified with horns or
protuberances in either sex. Elytral intervals alutaceous,
biseriately setigerously punctate-tuberculate. Readily
distinguished from all other small Florida species by
the maculate elytra. Similar to aciculatulus, but that
species is black, and the eyes are wider (about six
facets) dorsally.

TAXONOMIC NOTES: This is a distinctive species
not easily confused with any other. Florida specimens,
according to Howden and Cartwright (1963:86), have
the pronotal punctures larger and with the tubercles
often lacking, but in other respects they do not differ
from the remainder of the population.

It has been recorded from Connecticut south to Flor-
ida, west to eastern Texas, and north to Wisconsin.
However, it is not uniformly found in the intervening
areas. It is found throughout Florida, including at
least one record from the Keys (Monroe Co.).

BIOLOGY: This is probably the most common dung
beetle in Florida. It is very abundant in fresh cow
dung and has been recorded from several other types
of dung. It is also found in rotting fungi, other vege-
table substances, and occasionally in carrion. I have
taken specimens in several liquid baits, including malt,

00" '

proprionic acid, asafoetida, amyl acetate, and yeast. In
a single pint jar containing fermenting yeast which
had been left for two nights I collected over 700 speci-
mens of this species. It appears to be most abundant in
sandy areas. The immature stages are unknown.
SPECIMENS EXAMINED: Over 4,000 from 57 Florida
localities, representing 146 collection records (for
complete data see Appendix 21).
SELECTED REFERENCES: Blatchley, 1910:919-920;
1928:13; Boucomont, 1932:319; Brown, 1926:101;
1928a:26; Dillon and Dillon, 1961:515, P1. 49, Fig. 13;
Horn, 1875:140-141; Howden and Cartwright, 1963:85-
88; Fig. 9, 52-53; Knaus, 1926:264; Miller, 1954:376;
Schaeffer, 1914:298.

(Fig. 177)
The Florida record of this tribe is based an a
single specimen of Oniticellus cubiensis Lap., recorded
by Schwarz (in a manuscript) and cited by Blatchley
(1929:13). This old record is doubtful, and recent col-
lecting has produced no further specimens. However,
since it does occur in the Bahamas (Matthews, 1966),
I have included it here on the chance that it again
might be found in Florida.
The tribe is primarily Ethiopian and Oriental, with
the few American representatives displaying all the
characteristics of relict species (Halffter and Matthews,
1966:255). There are nine genera known from the
World, representing three subtribes: Oniticellina, in
which our single species is placed; Drepanocerina; and
The tribe is represented in the Western Hemisphere
by only six species in three genera as follows: Drepano-
cerus reconditus Matthews, a species recently described
from above 4,000 feet in the Blue Mountains of Jamai-
ca; Liatongus californicus (Horn), known from high
altitudes in Utah, California, and Oregon; Oniticellus
militaris (Cast.), found in Brazil, and possibly re-
cently introduced from the Old World (Pereira, per-
sonal communication); Liatongus monstrosus Bates from
Mexico; 0. rhinocerulus Bates from Mexico and possi-
bly not congeneric with Old World Oniticellus (Halffter,
personal communication); and O. cubiensis Lap. from
Cuba, Jamaica, Bahamas, and the doubtful record from
Key West, Florida.
The tribe was discussed in detail by Janssens
(1953), who proposed the genus Euoniticellus in which
the species cubiensis Lap. was placed. However, the
name has been treated as a subgenus of Oniticellus by
Balthasar (1963), Matthews (1966), and Halffter and
Matthews (1966). If subgenera are to be recognized
in the genus, its taxonomicc isolation" (Matthews,
1966) would probably merit a separate subgenus. Until
a reevaluation is made of all the members of the tribe,

I see little advantage in maintaining existing, or
describing new, subgeneric names.
The tribe is most similar superficially to the On-
thophagini from which it can be distinguished easily
by the eight-segmented rather than nine-segmented
antennae, and the scutellum small but visible. All other
Florida members of the subfamily have the antenna
nine-segmented and the scutellum hidden.

Genus ONITICELLUS Serville
(Fig. 177)
Oniticellus Serville 1828:356 (often cited as 1825; fide
Blackwelder, 1957:933, the citation should be 1828
for part 2:345-832).
Euoniticellus Janssens 1953:9, 41.
TYPE SPECIES: Scarabaeus cinctus Fabricius 1775
(designation not known, but cited by Matthews, 1966).

DIAGNOSIS: Similar in general facies to Onthophagus,
but differs by having eight-segmented rather than
nine-segmented antennae (the reverse being listed by
Arnett, 1962:412), and the scutellum is visible.
(see description of our single species, O. cubiensis

TAXONOMIC NOTES: Janssens (1953) divided this
genus into two: Oniticellus and Euoniticellus. However,
Balthasar (1963) relegates the latter to subgeneric
status. I agree with Matthews (1966), that if subgeneric
categories are maintained, O. cubiensis is sufficiently
distinct to occupy a position by itself. However, one of
the key features of the "genus" Euoniticellus is the
projecting genal margin, a character which is absent in
O. cubiensis.
record of this genus is based on a single specimen of
0. cubiensis from Key West recorded by Schwarz
(in a manuscript), and cited by Blatchley (1928:13).
Since this record is doubtful, and recent collecting has
produced no specimens, it probably does not occur
here now. However, it does occur in the Bahamas,
as well as Cuba and Jamaica (Matthews, 1966), and
I have treated it here so that it might be recognized
if it is found in Florida again. The genus is represent-
ed in the New World by this species and one in
The genus contains 23 species distributed as fol-
lows: Ethiopian (14), Palearctic (4), Oriental (3),
West Indian (1), and Mexican (1). Matthews (1966:
27) believed the genus was Ethiopian in origin and
probably entered North America from Asia via the
Bering Bridge. The relict distribution in the New
World parallels that in the genus Sisyphus, and I be-
lieve such patterns will require further zoogeographic
studies for logical explanation. Certainly the Bering
land bridge did permit the introduction of many
organisms into the New World. However, the dis-
tribution of one species in the West 'Indies and one

in Mexico, while the genus is lacking in the remainder
of North America and absent in colder climates, would
appear to substantiate the theory of continental drift.
The distributions of such relict species is of consider-
able academic interest, but there is little fossil evi-
dence to assist in arriving at an understanding
BIOLOGY: All of the species are apparently copro-
phagous, although a Mexican species in the related
genus Liatongus is associated with debris in the nests
of leaf-cutting ants (Atta sp.). None of the species has
been studied in any detail, but most of them appear
to deposit a single egg into a relatively small dung
mass at the end of burrows below the dung source. This
behavior is similar to that of many Onthophagus.
0. cubiensis occurs almost exclusively in open terrain
and was never found by Matthews (1966:30) in forests
or at the edges of wooded areas, and he found no
edaphic preferences.
SELECTED REFERENCES: Arnett, 1962; Blatchley,
1928; Halffter and Matthews, 1966; Janssens, 1953;
Matthews, 1966.

Oniticellus cubiensis Laporte
(Fig. 177)
Oniticellus cubiensis Laporte de Castelnau 1840:92.
Euoniticellus cubiensis (Lap.), Janssens 1953.
DIAGNOSIS: Typical Scarabaeinae, most similar to

S/ 'X i

Fig. 177. Oniticellus cubiensis

Lap., line = 3mm.

species of Onthophagus, from which it can be distin-
guished by the presence of a visible scutellum and
eight-segmented antennae. Length: 6.5-7.5 mm; width:
3-4 mm. Dull, light brown with extensive dark brown
markings over all surfaces, arranged in a pattern as
in Fig. 177; distinct cupreous reflections on head and
pronotum. Pygidium with a central dark spot. Clypeus
not dentate or emarginate; head entirely strongly mar-
gined, occipital carina strong and complete. Base of
pronotum not margined; surface very densely and
coarsely punctate with punctures of two sizes; median
longitudinal sulcus basal, rather deeply impressed.
Elytra leaving edges of abdominal sternites uncov-
ered laterally; eight shallow striae, indistinctly punc-
tate; glabrous except for a few long setae emerging
in an irregular row from first interstria very near
posterior apex.
Male with a short, stout horn on middle vertex,
a transverse ridge across middle of clypeus, followed
by a pair of curved ridges on clypeus between clypeo-
genal angles and horn. Fore tarsus with first segment
globular, inserted near base of fore spur. Female with-
out clypeal ridges or horn, with only one wide, straight,
transverse clypeo-frontal carina. Fore tarsus with first
segment long, linear, inserted well away from spur.

TAXONOMIC NOTES: The species appears to occupy
an isolated taxonomic position within the genus, one
which led Matthews (1966) to suggest that it belongs
in a subgenus of its own, if subgeneric categories are
maintained. It is distinguished from other members
of the genus by the shape of the head, a head horn
in the male, prosternal tumescence confined to the
female, and especially the absence of a distinct trans-
verse row of long hairs on the distal edges of the elytra.
from Jamaica, Cuba, Isle of Pines, and New Providence
and Eleuthera in the Bahamas. Blatchley (1928:13)
stated that it was ". . . recorded by Schwarz (MS.)
as having been taken at Key West by Morrison." This
is the only record from Florida. I have searched (with-
out success) for specimens at the U. S. National
Museum to substantiate this record. The manu-
script referred to by Blatchley is an annotated copy
of Schwarz's list of Florida Coleoptera (1878) which
Blatchley said was in the Smithsonian library. I have
been unable to locate this manuscript.
Although there is some doubt about the Florida
record, I have included the species here to permit its
identification should additional specimens be found.
I have personally searched for it without success in
various kinds of dung on the Florida Keys. The Key
West area is so well developed that few natural areas
remain. The supply of animal dung is very low since
no cattle are kept there. There are presently only about
two horses and several dogs and cats to provide a dung
supply. Human dung is also in short supply since
"privies" are now illegal.
The West Indian distribution is rather interesting.
Matthews (1966) stated that it is the only member of
the subfamily Scarabaeinae in the Greater Antilles
which occurs on more than one island. He further

stated that ". . . the apparent absence of any geo-
graphical variation suggests that it recently jumped
from one island to the others." If this is true, its
presence in the Bahamas suggests possible future im-
migration into Florida.
BIOLOGY: It is a common species in cow dung in
Cuba and Jamaica, but is never found in forested
areas (Matthews, 1966:30). I have collected a large
series (50+) along a cleared clay road through pine
forests in the province of Pinar del Rio, Cuba. There
appear to be no edaphic preferences (a fact which
would permit its establishment in the soil-poor Florida
Keys). Matthews found it in Jamaica at altitudes up
to 2,400 feet, but it was more abundant at sea level.
I have also found it common in many areas of Jamaica.
Its behavior has not been studied, and the imma-
ture stages have not been described. In most of the
species of Oniticellus the female lays a single egg in
a small mass of dung at the end of a burrow beneath
the source.
SPECIMENS EXAMINED: About 400, from Cuba and
SELECTED REFERENCES: Blatchley, 1928:13; Chev-
rolat, 1864:410; Gowdey, 1926:18; Gundlach, 1891:147;
Janssens, 1953; and Matthews, 1966:27-32, Fig.
14-18, 24.

(Fig. 178-309)
TYPE GENUS: Aphodius Illiger 1798 (by tau-
The higher categories of the Aphodiinae (i.e. tribes,
subtribes, groups, and genera) are poorly understood,
and nearly every reference has a different arrange-
ment. Part of the problem is due to the tremendous
numbers of species, at least some of which fill the
gaps between any arrangement of higher categories
devised. Until the subfamily receives additional study
on a world basis, the arrangement, even at the generic
level, will be very unsatisfactory.
Schmidt (1910a, 1910b) divided the subfamily into
five "groups": Aphodiina, Eupariina, Psammobiina,
Rhyparina, and Corythoderina. Balthasar (1964) rec-
ognized seven tribes: Aphodiini, Eupariini, Psammo-
biini, Rhyparini, Corythoderini, Thinorycterini, and
Demarziellini. Representatives of only the first three
are known from the U. S., and all three are repre-
sented in Florida.
Among the three tribes in Florida, there are genera
which have been placed in more than one tribe and
others which cannot be properly assigned. For this
reason I have not utilized the tribes in the treatment
which follows, although their current usage is shown
in the checklist in the introduction.

Specific examples of confusion about generic rela-
tionships are numerous, but I will list only a few here.
The genus Myrmecaphodius (Fig. 212) was recently
described for a single species found in the nests of
the imported fire ant (Solenopsis invicta Buren). It is
related to the genus Euparia, of which the single
species castanea (Fig. 220) occurs in Florida in the
nests of another ant (Solenopsis geminata). If we look
only at these two species there would be little doubt
about their generic distinction on the basis of numer-
ous gross morphological characters. However, if we look
at other species of Euparia from Central and South
America, it is not difficult to bridge these gaps. A
good example is E. vandykei, which could be placed in
either genus, depending upon the weight given to cer-
tain characters. In some cases the intermediate species
is described as another genus because it cannot be as-
signed easily. An example of the latter is Aphotaenius
which was described for a species which has charac-
ters of both Aphodius and Ataenius, and in both of
which it had been placed by previous authors. The
latter two genera are placed in separate tribes by all
authors. The genus Aphotaenius then must be arbi-
trarily assigned to one of these tribes. I maintain that
there is little value in using tribes at this stage in our
The latest catalogue (Schmidt, 1910b) listed 1,166
species for the world, but numerous species have been
described since that time. There are currently over
300 described species known from America north of
Mexico. The two largest genera, Aphodius and Atae-
nius, are both in need of revision. The latter is cur-
rently under study by 0. L. Cartwright of the U. S.
National Museum.
The subfamily is characterized as follows: clypeus
dilated to cover mandibles and other mouthparts (oc-
casionally maxillary palpi visible from above); anten-
nae nine-segmented, the club three-segmented; middle
coxae contiguous or nearly so; posterior tibiae with
two apical spurs; six visible abdominal segments; tarsi
with distinct claws; body shape always oblong, some-
what cylindrical.
The larvae are characterized as follows (Ritcher,
1966:26): antenna four- (or apparently five) segment-
ed with the last segment reduced in size, third segment
usually with apical process; epipharynx trilobed, pe-
dium surrounded by phobae; tormae united, with
prominent epitorma; galea and lacinia of maxilla dis-
tinctly separate, but often close together; maxillary
stridulatory teeth sometimes absent; dorsum of thoracic
and abdominal segments plicate; concavities of respira-
tory plates of thoracic spiracles facing posteriorly, those
of abdominal spiracles facing ventrally or cephaloven-
trally; legs four-segmented, with well developed claws;
and lobes whitish or yellowish, bare of setae. For
keys to the known U. S. larvae and additional descrip-
tions, see the paper by Jerath (1960b).
The biology and ecology of the subfamily is so
diverse that few generalizations can be made. Many
of the species feed on dung or decaying vegetable
matter. Some of the dung feeding species are known
only from the dung of one animal (e.g., aeer, Geomys,

Gopherus, etc.). Some are found only along the sea-
coast and others only at high altitudes. Several species
are myrmecophilous or termitophilous. Many of the
species are abundant at lights. The North American
literature contains little information about the ecology
of this group, except for a paper on the Oregon fauna
by Jerath and Ritcher (1959). However, there is an
excellent account of the ecology of European species,
with special reference to Sweden (Landin, 1961).

SELECTED REFERENCES: Arnett, 1962:412-414;
Balthasar, 1964:1-652; Blatchley, 1910:920-935; 1928:
13-14; 22-28; Cartwright, numerous papers 1934-1965;
Dillon and Dillon, 1961:516-523; Horn, 1870b: 110-
134; 1871a: 284-297; 1887:1-110; Jerath, 1960b:43-94
(larvae); Jerath and Ritcher, 1959:169-175; Landin,
1961:1-228; Mohr, 1930:263-284; Ritcher, 1958:311-
334; 1966:26-29 (larvae); Schmidt, 1910a:1-155;
1910b:1-111; 1912:1-11; 1922:1-614.

Key to the Florida genera of Aphodiinae

1. Pronotum usually with transverse swellings and
grooves; median longitudinal impression usually
present; head rugulose to verrucose; clypeus
often dentate; posterior tarsal segments usually
short, triangular; posterior tibial spurs often
twisted, flattened on one side, and spatulate-
shaped ................................ . 7
1'. Pronotum without transverse swellings and
grooves, median longitudinal impression rarely
evident; head usually without rugulae, although
often with feeble vermiculate lines; clypeus den-
tate or not; posterior tarsal segments usually
elongate, not expanded or triangular; posterior
tibial spurs usually long, narrow and acute.. .2
2(1'). Middle and posterior tibiae with transverse
carinae; head often tuberculate; some species
bicolored or maculate ..................... 3
2'. Posterior tibiae without transverse carinae, al-
though there is often at least one on the middle
tibiae; head never tuberculate; color brown to
black, never bicolored or maculate.......... 4
3(2). Middle and posterior tibiae with a terminal
fringe of setae or spicules ..................
(Fig. 178) ....................... Aphodius
3'. Middle and posterior tibiae with the terminal
fringe replaced by two small, triangular teeth,
each with a fine, hairlike seta basally on each
side. (Fig. 292) ............................
.............. Aphotaenius carolinus (Van D.)
4(2'). Pronotum explanate or expanded at the sides;
posterior tibia curved inwardly, slightly expand-
ed at the apex; myrmecophilous ............ 5
4'. Pronotum not explanate at the sides; posterior
tibia straight or feebly curved, not expanded at
the apex; not myrmecophilous ...........6...

5(4). Elytra deeply notched at the base inside the
humeri which are prolonged into a cuneiform
process; lateral pronotal margin with fine, non-
contiguous setae; dorsal surface covered with
fine yellowish pubescence, the setae arising from
tuberculate punctures; genae separated from
clypeus by a deep notch; pygidium smooth, not
verrucose. (Fig. 220) ....................
. ................. Euparia castanea Serville
5'. Elytra not deeply notched at the base, the
humeri dentate but not prolonged; lateral pro-
notal margin with flattened, contiguous, blunt
setae; dorsal surface glabrous; genae feebly sep-
arated from the clypeus; pygidium verrucose.
(Fig. 212) ................................
..... Myrmecaphodius excavaticollis (Blanch.)

6(4'). Mesosternum not carinate, clypeus without teeth;
terminal segment of maxillary palpi slender,
elongate; middle and posterior tarsi one-third
longer than their tibiae; sexual dimorphism
noticeable; male with anterior tibiae bidentate,
the apical spur long, twisted, and incurved at
the tip; female with anterior tibiae tridentate,
apical spur short, straight, and acute. (Fig. 290)
.............. Pseudataenius socialis (Horn)
6'. Mesosternum carinate; clypeus sometimes den-
tate; terminal segment of maxillary palpi thicker
at the middle and posterior tarsi never as much
as one-third longer than their tibiae; sexual
dimorphism not noticeable, the male never with
anterior tibial spur twisted and incurved at the
apex. (Fig. 222) .................. .Ataenius

7(1). Long spur of posterior tibia at least as long as
first two tarsal segments; tarsus distinctly shorter
than tibia; middle and posterior tarsal segments,
especially the first one, expanded at the apex,
somewhat triangular. (Fig. 293). . .Psammodius
7'. Long spur of posterior tibia usually shorter,
rarely longer, than the first tarsal segment;
tarsus as long as tibia; middle and posterior
tarsal segments not greatly expanded .... ...8

8(7'). Pronotum without lateral fringe of setae; small-
est species of Scarabaeidae (length 1.9-3 mm);
pygidium with 6-10 elongate setae; elytral in-
tervals convex, not carinate or costate. (Fig.
306) ....................... Pleurophorus
8'. Pronotum margined laterally with a fringe of
spatulate setae; larger (length 3-4 mm); pygi-
dium without elongate setae; elytral intervals
costate. (Fig. 309) .... Rhyssemus scaber Hald.

Genus APHODIUS Illiger
(Fig. 178)
Aphodius Illiger 1798:15.

TYPE SPECIES: Scarabaeus fossor Linnaeus 1758 (by
subsequent designation of Curtis, 1824).
DIAGNOSIS: Mandibles concealed beneath the clypeus;
pronotum without transverse swellings and impressions
or rows of punctures; head never granulate or verru-
cose; tarsal segments cylindrical, elongate, never tri-
angular; middle and posterior tibiae with transverse
carinae (Fig. 41); elytral intervals never carinate,
simply convex or flattened; several species pale yellow,
and a few bicolored with red and black; length 2 to
8 mm; width 1 to 4 mm.
TAXONOMIC NOTES: The genus was established in
1798 to include 32 species, but no type species was
designated. Curtis' designation cited above was ac-
cepted by Chapin (1940), and the subsequent type
designations of Scarabaeus oblongus Scopoli by Gozis
(1886) and Scarabaeus erraticus Linnaeus by Paulian
(1935) are both invalid.
Schmidt (1922a) listed 74 subgenera of Aphodius.
Some of these names also have been elevated to generic
rank, especially by European authors. The genus is so
large (with 1,000+ species described) that a world-
wide revision is not imminent.
It has a nearly cosmopolitan distribution, and sev-
eral species have been transported accidentally around
the world by man. Until a thorough study of the
world fauna is made, it is futile to divide the species
into natural groupings. The U. S. species were placed
in groups with letter designations by Horn (1887),
but these are of little value at the present time. I
have therefore used Aphodius in its broadest sense
without reference to subgeneric names, except to point
out where the species were placed by Schmidt.
The problem of subgenus in the large genus
Aphodius was discussed by Grebenscikov (1956). He



Fig. 178. Aphodius fimetarius (L.), line = 5mm.

believed that the evolutionary unit should be either
a genus or not, and that the concept of subgenus was
not tenable. My personal belief is that the use of
subgenera or species groups permits organization within
large genera (e.g., Aphodius and Phyllophaga) without
the attendant nomenclatural problems associated with
generic names (and the binomial).
The U. S. species are in need of revision, although
the Florida fauna is fairly well-known. The latest revi-
sion of the U. S. species is by Horn (1887), and the
number of species has been more than doubled since
that time. Few subspecies of U. S. species have been
described, and most of these are of doubtful validity.
The genus is the only representative of the tribe
Aphodiini in Florida. Its nearest relative in our fauna
is the genus Aphotaenius which is presently placed in
the Psammodiini.
nearly world-wide in distribution with about 1,000
described species. Nearly 200 species are known from
the U. S., 17 of which are here reported from Florida,
with two others doubtfully recorded. Five of the Flor-
ida species have been introduced from the Old World
and are now widely distributed in the New World;
this is especially true for A. lividus which is recorded
from nearly all parts of the globe. Many of these in-
troduced species are more abundant than our native
ones and are more plentiful in their new territories
than in their native habitats. Most of these introduc-
tions occurred more than 50 years ago, and the beetles
were most likely transported in ship ballast or in slave
and cattle boats where dung was readily available.
Only two of the Florida species (A. lividus and
A. cuniculus) reach the West Indies, and both are
widespread in the Caribbean and Gulf area. Only one
West Indian species (of the four known from there)
has been reported as endemic (from Cuba).
Apparently several of the introduced species are
continuing to expand their range. One such example
is A. haemorrhoidalis, which was not found in Florida
prior to 1967, although it is common in cow dung
throughout the eastern U. S. Only one species (A.
floridanus) is truly endemic in Florida, although those
found in the burrows of the gopher tortoise and the
pocket gopher are nearly endemic.
BIOLOGY: Nearly all members of the genus feed on
dung of the higher animals. Several species are known
only from the burrows of rodents (e.g., Geomys,
Thomomys), and one is associated with the gopher
tortoise (Gopherus polyphemi Daudin). Other species
have been found in the dung of deer, pack rats, fox,
etc., but are particularly abundant in cow dung. All
of the Florida species (except parcus) are found in
dung. A. parcus is sometimes collected in large num-
bers at light, especially in coastal localities, but it has
not been found feeding, or even hiding, in dung.
Although this group probably contains as much
diversity in habits as the other primary dung feeders
(Scarabaeinae), they have not been as thoroughly
studied. One exception is the recent work by Landin
(1961 and 1968) in which the ecoldgy of several

European Aphodius is detailed. He concluded that the
factors governing the distribution of the species are
mainly of a climatic (and microclimatic)
McDaniel and Balsbaugh (1968) found two species
of Aphodius overwintering in frozen cow dung in South
Dakota. Many species of Aphodius have been found
on snow in the winter, and some species fly in swarms
(emerging from hibernation) during early spring in
northern latitudes. There are also many alpine species
in the Palearctic region, and I have collected them in
the high paramo in South America.
A few species have been reported as pests of culti-
vated mushrooms, and at least one species (A. pardalis)
is a turf pest in the Pacific northwest. The following
Florida species are attracted to light: aegrotus, cam-
pestris, cuniculus, floridanus, laevigatus, lividus, parcus,
rubeolus, and troglodytes. Certain species are attracted
to artificial baits such as malt; among which are the
following Florida species: campestris, cuniculus, and
stupidus. Some species (e.g., stupidus, bicolor, and
crassulus) are active primarily in the winter or early
spring. Others, such as campestris, fimetarius, and
lividus, are active nearly the entire year, especially
in the southern half of the state. Some of the species
are extremely abundant and can be found by the hun-
dreds in a single cow dropping. Landin' (1961:208)
reported that the competition within a single dropping
was individual rather than interspecific or intraspecific.
He concluded that the fluctuations of populations de-
pended on " . . . . abiotic environmental factors rather
than on the competition factor." The larvae of 16
U. S. species were described by Jerath (1960b).
SELECTED REFERENCES: Balthasar, 1964; Blatch-
ley, 1928; Brown, 1927b; Chapin, 1940; Grebenscikov,
1956; Horn, 1870; 1887; Jerath, 1960b; Landin, 1961;
1968; Schmidt, 1910a; 1910b; 1922a.

Alphabetical listing of Florida Aphodins with the
subgenera In which they were placed by
Schmidt (1922a)
aegrotus Horn (not placed, but would fit Koshantschi-
bicolor Say (Cinacanthus)
campestris Blatchley (not placed, but would fit
crassulus Horn (Mendidius)
cuniculus Chev. (Nialus); the synonym vestiarius Horn
was placed in the subgenus Oromus
fimetarius (L.) (Aphodius)
floridanus Rob. (not placed, but would fit Agrilinus)
granarius (L.) (Calamosternus)
haemorrhoidalis (L.) (Teuchestes)
haldemani Horn (Platyderides)
laevigatus Hald. (Koshantschikovius)
lentus Horn (Amidorus)
lividus (Oliv.) (Nialus)
lutulentus Hald. (Amidorus)

parcus Horn (not placed by Schmidt, but placed in
the genus Didactylia by Brown, 1929a).
rubeolus Beauv. (Koshantschikovius)
rusicola Melsh. (Agrilinus)
stercorosus Melsh. (Koshantschikovius)
stupidus Horn (Amidorus)
troglodytes Hubbard (Koshantschikovius)

Key to the Florida species of Aphodius
1. Surface of elytra dull, alutaceous, opaque;
pronotum densely punctate .............. 2
1'. Surface of elytra shining, never alutaceous;
pronotal punctation variable from dense to
impunctate ............................. 4

2(1). Mesosternum carinate; color red-brown;
length about 3.5 mm; Florida record (Pensa-
cola) doubtful................. lentus Horn
2'. Mesosternum not carinate; color dull gray to
brown-black; length 3-7 mm............. 3

3(2'). Clypeus on each side rounded, barely angulate;
elytral pubescence obvious, evenly distributed;
length 3-5 mm; common in northern Florida,
especially in winter ........... stupidus Horn
3'. Clypeus on each side angulate; elytral pubes-
cence short, inconspicuous, scattered; length
5.5-7 mm; only two Florida records........
........................... lutulentus Hald.

4(1'). Size larger (length 6-8 mm; width 3-4 mm);
elytra red or brown, never black..........5
4'. Size smaller (length 2-6.5 mm; width 1-2.5
mm); elytral color variable from black to pale
yellow ............................... 6

5(4). Bicolored, elytra red, the pronotum black with
anterior angles red. Head with three promi-
nent tubercles; common species in cow dung
......... ............... fimetarius (L.)
5'. Color uniform red-brown; head with trans-
verse carina, but no prominent tubercles; oc-
curring in burrows of the Florida pocket
gopher (Ceomys) and rarely at light..... 19

6(4'). Head tuberculate, at least with three convex
areas on the frons, ground color mainly black
(except lividus which is pale with smoky
markings) ............................ 7
6'. Head not tuberculate, never with more than
the frons noticeably convex; ground color
brown to yellow (dorsally black and ventrally
orange to reddish in bicolor) ............ 13

7(6). Clypeus with two prominent teeth; primarily
a winter species of the northern half of the
state ....................... crassulus Lee.
7'. Clypeus without prominent teeth; not pri-
marily winter species.................... 8

8(7'). Elytra tipped with a broad red area; scutel-
lum nearly twice as long as broad..........
...................... haemorrhoidalis (L.)
8'. Elytra without red spots at the tip; scutellum
only slightly longer than broad, often barely
punctate to impunctate................... 9

9(8'). Base color of elytra and pronotum yellow
with smoky brown markings... lividus (Oliv.)
9'. Base color black to dark red brown, nearly
unicolorous .......................... 10

10(9'). Basal marginal line of pronotum absent;
smaller species (length about 3 mm); com-
mon throughout Florida at light............
........................... cuniculus Chev.
10'. Basal marginal line of pronotum complete;
larger species (length 4-5 mm); rarely col-
lected at light ........................ . 11

11 (10'). Scutellum depressed, surrounded by a depres-
sion; pronotal punctures widely and unevenly
scattered; first segment of posterior tarsi
shorter than the long spur; a single Florida
record (Jackson Co.)......... granarius (L.)
11'. Scutellum convex, not surrounded by a depres-
sion; pronotal punctures more evenly distrib-
uted and denser; first segment of posterior
tarsi as long or longer than the long tibial
spur ................................. 12

12(11'). Elytral intervals convex; pronotal punctures
larger, denser, and of two distinct sizes; found
sparingly the length of the Florida peninsula
........................... floridanus Rob.
12'. Elytral intervals more flatterned; pronotal
punctures smaller, less dense, and nearly uni-
form in size; Florida record doubtful........
........................... rusicola M elsh.

13(6'). Dorsal surface black, venter red to orange;
only three Florida records; larger (length
5.5-6.5 mm) .................. bicolor Say
13'. Nearly uniform red brown to pale yellow;
smaller (length 2-5 mm) ............... 14

14(13'). Lateral pronotal margin with setae.......15
14'. Lateral pronotal margin without setae..... 18

15(14'). Setae of lateral pronotal margin elongate;
small (length 2-3 mm); pale yellow, often
with smoky markings on the pronotum; trans-
verse carinae of middle and posterior tibiae
obsolete ....................... parcus Horn
15'. Setae of lateral pronotal margin short; larger
(length 3-4.5 mm); usually red-brown, at least
the pronotum; transverse carinae of middle
and posterior tibiae well developed.......16

16(15'). First segment of posterior tarsi slender, elon-
gate, longer than the long tibial spur; none
of elytral intervals pubescent; found in the
burrows of the Florida gopher tortoise
(Gopherus polyphemi) ... troglodytes Hubbard
16'. First segment of posterior tarsi shorter or
equal to the long spur; elytra pubescent on
lateral posterior one-third; found in cow dung
and at light............................17

17(16'). Spurs of posterior tibia short, spatulate, with
a broad concave surface; first posterior tarsal
segment shorter than the long tibial spur;
color nearly uniformly red-brown...........
........................ rubeolus (Beauv.)
17'. Spurs of posterior tibia elongate, narrow, with
a narrow concave surface; first posterior tarsal
segment about equal to the long tibial spur;
color somewhat variable, the elytra often paler
yellow than the pronotum, rarely red-brown
throughout ............. campestris Blatch.

18(14'). Basal marginal line of pronotum complete;
short and broad (width 2 mm); pronotal punc-
tures coarse at lateral one-third, the disc with
very minute punctures; tarsal claws elongate
(nearly two-thirds the length of last tarsal
segment); found in burrows of the pocket
gopher (Geomys spp.) and at light.........
............................ aegrotus Horn
18'. Basal marginal line of pronotum absent; elon-
gate narrow (width 1.5 mm); at least some of
the coarse pronotal punctures encroaching on
the disc; tarsal claws normal (about half the
length of the last tarsal segment); Florida
record doubtful, common in cow dung in
N. E. ................... stercorosus Melsh.

19(5'). Male anterior tibial spur broad and spatulate
shaped (Fig. 180); pronotum explanate at
sides; a single Florida record.............
......................... haldemani Horn
19'. Anterior tibial spur normal, narrow and point.
ed (Fig. 179); pronotum normal, not explanate
at sides; frequent locally at light ..........
.......................... laevigatus Hald.

Aphodius aegrotus Horn
Aphodius aegrotus Horn 1870b:127-128.
Aphodius geomysi Cartwright 1939b:356-357. (NEW
DIAGNOSIS: Dorsal surface glabrous, shining, deep
red-brown, broad, and short, elliptical in outline. Head
without tubercles, lightly punctate with fine punctures.
Pronotum extremely finely punctate throughout and
with a group of coarse punctures on the lateral one-
third, some of which are elongate in, shape; lateral

Fig. 179-180. Anterior tibiae and tarsi of Aphodius spp.
(males): 179) A. laevigatus Hald., 180) A. haldemani Horn.
Note shape of tibial spurs.

margin without fringe of setae; basal marginal line
complete. Elytra without any trace of pubescence,
finely punctate. Tarsal claws narrow and extremely
elongate for the genus.
Similar in color to rubeolus from which it can be
separated by the elongate, acute posterior tibial spurs.
Of the Florida species, it appears most closely related
to troglodytes from which it can be separated by the
lack of marginal setae on the pronotal sides. Of the
U. S. species, it is most closely related to cavidomus
Brown, known from prairie dog burrows in Oklahoma,
from which it can be separated by its larger size,
greater convexity, and elliptical shape.
TAXONOMIC NOTES: This species was described by
Horn from "North Carolina" and was subsequently
reported by him (1887) from Florida. Nothing has
been published on the species since Horn's works, and
it has long been unrecognized. The type specimen is
apparently lost, since it was not found at the Phila-
delphia Academy of Sciences with the remainder of
Horn's collection. I have examined the three speci-
mens labeled as aegrotus in the Horn collection, and
all are from Florida. They all carry the accession
number 5549 of the Horn collection. One of these
is A. troglodytes Hubbard, and was probably a cura-
torial misplacement rather than a misidentification
by Horn. The other two are presumably the basis of
Horn's record for Florida. One bears the "Fla." label
and the other '"Marion County" without reference to
a state. These specimens are conspecific with paratypes
of A. geomysi Cartwright (1939b), which is the basis
of the above synonymy. It is unfortunate that this
latter name requires suppression, since it indicates the
relationship with the pocket gopher.
It was described from "North Carolina," but no further
specimens have been seen from there. It is known

otherwise only from Florida where it is recorded from
Pensacola in the western panhandle to northern High-
lands County in the south. Its range is probably co-
extensive with its host (Geomys pinetis and related
species of the southeast (Fig. 75). Blatchley (1929)
had not examined any Florida specimens and merely
repeated Horn's records. However, I found two speci-
mens in the Blatchley collection at Purdue University
from Dunedin, Florida, but they were erroneously de-
termined as rubeolus. The type locality of geomysi is
Newnan's Lake, Alachua County, Florida. Other local-
ities mentioned in the original description of that
species include: Gainesville, Eau Gallie, Lake Streaty,
and Marianna. Hubbell and Goff (1939:161) also
reported it [as geomysil from Melrose and Welaka.
I have seen it from 10 additional localities.
BIOLOGY: As mentioned earlier this species is found
in the burrows of the Florida pocket gopher (Geomys
pinetis ssp.), where it feeds on dung. It is often very
common in this habitat, but has never been found in
any other kind of dung. It is not uncommon at light,
but is never abundant. Practically nothing is known
of the habits within the burrows. The tarsal claws are
more elongate than in other species, a situation which
might provide some freedom from dislodgment when
the host creates a disturbance. Specimens have been
collected every month except December and January.
The immature stages are unknown.
SPECIMENS EXAMINED: 196 from 17 Florida local-
ities, representing 87 collection records (for complete
data see Appendix 22).
SELECTED REFERENCES: Blatchley, 1928:24; Cart-

wright, 1939b:356, 357; Horn, 1870b:127-218; 1887:43;
Hubbell and Goff, 1939:161; Schmidt 1922a:333-334.

Aphodius bicolor Say
Aphodius bicolor Say 1823:212.
Aphodius comanchi Robinson 1940:146.
DIAGNOSIS: Distinguished from nearly all other
species by the black upper surface and the red to
orange ventral surface. Clypeal angles each side of
the emargination prominent and reflexed. Head, pro-
notum, and elytra densely, fairly coarsely punctate.
Tips of the elytra with an alutaceous patch, but other-
wise glabrous. Pronotal midline often impunctate, shin-
ing; basal marginal line complete. Legs orange-red
like the venter; first segment of anterior tarsi much
shorter than the second; first segment of posterior tarsi
longer than the long tibial spur. Rare in Florida and
found primarily in the winter (September - April).
TAXONOMIC NOTES: A. comanchi Robinson was
described from Texas, but later synonymized by its
author (1947:150) as only a larger specimen of bicolor
Say. The species is variable in length (4.5 to 6.5 mm)
and color. Horn (1870b:130) mentioned specimens
which had the elytra "dark ferrugineus with the alter-
nate intervals (2-4-6) showing paler rounded spots."
I have not seen this color variation, and all the Florida
specimens have the elytra entirely black.
There is considerable sexual dimorphism in the
anterior tibiae and the posterior femora. In the male
the anterior tibial spur arises almost opposite the lower
tooth, a position rare within the genus; posterior femur
fimbriate on the posterior margin, as well as on the
inner margin of the posterior tibia. In the female the
anterior tibial spur arises almost opposite the middle
tooth, a normal position in the genus; posterior femur
has only a few setae on the posterior margin, but is
not fimbriate, nor is the inner face of the posterior
It is fairly widely distributed in the U. S. although
rarely abundant. It has been recorded from the fol-
lowing states: Alabama (Loding, 1933), Florida
(Blatchley, 1919), Illinois (Mohr, 1943), Indiana
(Blatchley, 1910). Louisiana (Summers, 1874). Massa-
chusetts (Horn, 1887), Missouri (Say, 1823), Nebraska
(Dawson, 1922), New Jersey (Smith, 1910), New York
(Leng, 1928), North Carolina (Brimley, 1938), Ohio
(Dury, 1902), Oklahoma (Brown, 1928a, South Caro-
lina (Cartwright, 1934b), Texas (Horn, 1870b), and
Canada (Wickham, 1894).
In Florida it has been collected only three times,
from Dunedin on the west coast to Macclenny in the
northeast (see section on specimens examined for
complete data).
BIOLOGY: Very little has been published on this
species. It is primarily a winter species (September-

April), but is rarely abundant. It has been found on
a wide variety of soil types and appears to be more
common in wooded situations. Mohr (1943) reported
taking 123 specimens in Illinois from a single pile of
cow dung which was deposited either the night be-
fore or early that morning. I collected nearly 100
specimens in Ohio in a pastured woodlot in September.
The single Florida specimen that I have collected
was taken in a malt trap set in a farm woodlot, heavily
grazed by cattle and with no undergrowth. Several
hours of collecting at this locality in cow dung, of
various ages, failed to produce additional specimens.
The other Florida specimen, on which the habitat was
recorded, was in a wooded area in the dung of a wild
animal, possibly a wild pig. The immature stages
are unknown.
SPECIMENS EXAMINED: Approximately 115, only
3 of which are from Florida as follows: (1) Baker Co.,
Macclenny, 9-XII-60, R. E. Woodruff and E. W. Hold-
er, Jr., malt bait trap; (1) Levy Co., 10 mi. N. of
Cedar Key, 7-1-67, A. L. O'Berry, pig (?) dung; (1)
Pinellas Co., Dunedin, 18-III-18, W. S. Blatchley [PUI.
SELECTED REFERENCES: Blatchley, 1910:933;
1919:31; 1928:24; Dillon and Dillon, 1961:520, P1. 50,
Fig. 8; Horn, 1870b:130; 1887:46; Mohr, 1943: 292;
Robinson, 1947a:150.

Aphodius campestris Blatchley
Aphodius campestris Blatchley 1912:330.

Aphodius compestris Blatch., Loding 1945:99 (misspel-

Aphodius stercorosus Melsh., Frost 1964:142 (?misiden-

DIAGNOSIS: Small (length 2.5-4 mm), pale yellow
to yellow-brown, rarely red-brown, the pronotum often
darker than the elytra. Similar to rubeolus, from
which it is easly separated by the longer more narrow
posterior tibial spurs; first posterior tarsal segment
as long as the next three. In rubeolus the posterior
tibial spurs are short, broad, and spatulate; first poster-
ior tarsal segment distinctly shorter than the next
three. Also similar to stercorosus, which has not de-
finitely been recorded from Florida, but in that species
the basal marginal line of the pronotum is absent.
There is usually a patch of pubescence on the pos-
terior one-third of the elytra of campestris

TAXONOMIC NOTES: Although this species is fairly
distinctive, it has been confused with stercorosus in
collections, and it was probably the basis for the Flor-
ida record of that species by Frost (1964). Variation
exists primarily in size and color. The smaller speci-
mens (2.5-3 mm) are often much lighter colored than
the larger ones (4 mm). It is only these larger ones
which are reddish-brown and most similar in color to
rubeolus. Although not placed in a subgenus by
Schmidt (1922a), it would fall in his Koshantschiko-
vius. Brown (1927b) placed it in Horn's group "I-B"
along with rubeolus, stercorosus, lentus, tenuistriatus,
troglodytes, cynomysi, and cavidomus.

It was described originally from near Sarasota, Flor-
ida, and subsequently recorded from the following
states: Alabama (Loding, 1945), Georgia (Miller,
1954), New Jersey (Sim, 1930), North Carolina (Brim-
ley, 1938), and South Carolina (Cartwright, 1934b). It
probably occurs more widely in the eastern U. S., es-
pecially in the Gulf states and in sandy areas of the
east coast. In Florida it has been found in nearly all
areas of the state and probably occurs throughout.
It has not yet been collected in southern Dade or
Monroe Counties or in the Keys. It is interesting that
Blatchley (1928) recorded it only from three locations:
Sarasota, Dunedin, and LaGrange. Since it is presently
common throughout the year in cow dung and at light,
it is difficult to explain the paucity of early records,
unless it has greatly increased in abundance during the
past 40 years.

BIOLOGY: Although this is one of the most common
dung beetles in Florida, very little information is avail-
able on its habits. It is common in cow dung and also
has been found on deer, horse, and human dung. It
is one of the most commonly collected Aphodius in light
traps in Florida. I have collected large numbers in
malt bait traps, especially those to which a few drops
of proprionic acid have been added. A few specimens
were taken in rotting fungi, and some were collected
in a Berlese funnel sample of sand pine litter. Miller


(1954) collected specimens in traps baited with human
dung only between 5 P.M. and 10 A.M. None were
taken in traps between 8 A.M. and 7 P.M. Although
adults are not rare in cow dung during the daytime,
these data, along with the abundance at light, suggest
a nocturnal flight period. It appears to be more abund-
ant in very sandy soil conditions and was found in
the Pine Barrens of New Jersey under these conditions.
Adults have been collected every month of the year
in Florida. The immature stages are unknown.
SPECIMENS EXAMINED: Over 1,600 Florida speci-
mens from 80 localities, representing 384 collection
records (for complete data see Appendix 23), including
the type (Blatchley Collection, Purdue University).
SELECTED REFERENCES: Blatchley, 1914:91;
1928:23; Brown, 1927b:163-165; Miller, 1954:380,
Table 1; Sim, 1930:140, 142.

Aphodius crassulus Horn

Aphodius crassulus Horn 1870b:118.
DIAGNOSIS: The only Florida Aphodius with two dis-
tinct teeth on the clypeus. Black, shining, extremely
convex, compact, length 3.6 to 6 mm. Scutellum small;
posterior tibiae fringed with equal spinules; anterior
tibial face smooth, impunctate; first segment of anter-
ior tarsi shorter than second; head trituberculate. Cly-
peus with a few granules and punctaterrugose near the
margin. In general appearance it is similar to flori-

danus Robinson, from which it can be distinguished by
the bidentate clypeus, the more prominent and acute
genal angles, dark black color, and the much more
convex elytral intervals on the apical one-third.

TAXONOMIC NOTES: This species was placed by
Horn (1887) in his group "B" although he admitted
that this group was ". . . not very homogenous." Cart-
wright recently (1957) studied the group of species
closely related to crassulus, including the following:
bottimeri Cartwr., odocoilis Rob., brimleyi Cartwr.,
spiniclypeus Hinton, windsori Cartwr., lodingi Cartwr.,
abusus Fall, crassuloides Fall, and pseudabusus Cartwr.
It was placed in the subgenus Mendidius by Schmidt
A. crassulus is particularly variable in size (length
3.6 to 6 mm, although no Florida specimens have been
seen over 5 mm long). Sexual dimorphism is most
noticeable in the anterior tibial teeth and spur. In the
male the third or basal tooth is about normal in size,
but in the female it is reduced, making the middle
tooth appear much more elongate. The anterior tibial
spur is stouter and more curved in the male. The
clypeal teeth are much more prominent and reflexed
in the female.
It was recorded by Horn (1870b) from Georgia and
Florida, and in 1887 he listed it from "Florida to Tex-
as." It is also recorded from North Carolina (Brim-
ley, 1938), South Carolina, Virginia, and Georgia
(Cartwright, 1957). Records from Ohio (Dury, 1902),
New Jersey (Sim, 1930), and New Mexico (Fall and
Cockerell, 1907) probably refer to other species.


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