• TABLE OF CONTENTS
HIDE
 Title Page
 Preface
 Acknowledgement
 Table of Contents
 List of Figures
 Abstract
 Introduction
 General account of the family
 Preliminary checklist of florida...
 Systematic account
 Illustrations
 Appendix
 Bibliography
 Biographical sketch
 Copyright














Group Title: scarab beetles of Florida (Coleoptera: Scarabaeidae) Part I. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotru
Title: scarab beetles of Florida (Coleoptera: Scarabaeidae) Part I. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae)
CITATION PDF VIEWER THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00091315/00001
 Material Information
Title: scarab beetles of Florida (Coleoptera: Scarabaeidae) Part I. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae)
Series Title: scarab beetles of Florida (Coleoptera: Scarabaeidae) Part I. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotru
Physical Description: Book
Creator: Woodruff, Robert E.
 Record Information
Bibliographic ID: UF00091315
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000430506
oclc - 37545706

Downloads

This item has the following downloads:

UF00091315 ( PDF )


Table of Contents
    Title Page
        Page i
    Preface
        Page ii
        Page iii
        Page iv
    Acknowledgement
        Page v
        Page vi
        Page vii
    Table of Contents
        Page viii
        Page ix
        Page x
    List of Figures
        Page xi
        Page xii
        Page xiii
        Page xiv
        Page xv
        Page xvi
    Abstract
        Page xvii
        Page xviii
    Introduction
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
    General account of the family
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
    Preliminary checklist of florida scarabaeidae
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 42
        Page 43
    Systematic account
        Page 44
        Page 45
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
        Page 57
        Page 58
        Page 59
        Page 60
        Page 61
        Page 62
        Page 63
        Page 64
        Page 65
        Page 66
        Page 67
        Page 68
        Page 69
        Page 70
        Page 71
        Page 72
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
        Page 78
        Page 79
        Page 80
        Page 81
        Page 82
        Page 83
        Page 84
        Page 85
        Page 86
        Page 87
        Page 88
        Page 89
        Page 90
        Page 91
        Page 92
        Page 93
        Page 94
        Page 95
        Page 96
        Page 97
        Page 98
        Page 99
        Page 100
        Page 101
        Page 102
        Page 103
        Page 104
        Page 105
        Page 106
        Page 107
        Page 108
        Page 109
        Page 110
        Page 111
        Page 112
        Page 113
        Page 114
        Page 115
        Page 116
        Page 117
        Page 118
        Page 119
        Page 120
        Page 121
        Page 122
        Page 123
        Page 124
        Page 125
        Page 126
        Page 127
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
        Page 134
        Page 135
        Page 136
        Page 137
        Page 138
        Page 139
        Page 140
        Page 141
        Page 142
        Page 143
        Page 144
        Page 145
        Page 146
        Page 147
        Page 148
        Page 149
        Page 150
        Page 151
        Page 152
        Page 153
        Page 154
        Page 155
        Page 156
        Page 157
        Page 158
        Page 159
        Page 160
        Page 161
        Page 162
        Page 163
        Page 164
        Page 165
        Page 166
        Page 167
        Page 168
        Page 169
        Page 170
        Page 171
        Page 172
        Page 173
        Page 174
        Page 175
        Page 176
        Page 177
        Page 178
        Page 179
        Page 180
        Page 181
        Page 182
        Page 183
        Page 184
        Page 185
        Page 186
        Page 187
        Page 188
        Page 189
        Page 190
        Page 191
        Page 192
        Page 193
        Page 194
        Page 195
        Page 196
        Page 197
        Page 198
        Page 199
        Page 200
        Page 201
        Page 202
        Page 203
        Page 204
        Page 205
        Page 206
        Page 207
        Page 208
        Page 209
        Page 210
        Page 211
        Page 212
        Page 213
        Page 214
        Page 215
        Page 216
        Page 217
        Page 218
        Page 219
        Page 220
        Page 221
        Page 222
        Page 223
        Page 224
        Page 225
        Page 226
        Page 227
        Page 228
        Page 229
        Page 230
        Page 231
        Page 232
        Page 233
        Page 234
        Page 235
        Page 236
        Page 237
        Page 238
        Page 239
        Page 240
        Page 241
        Page 242
        Page 243
        Page 244
        Page 245
        Page 246
        Page 247
        Page 248
        Page 249
        Page 250
        Page 251
        Page 252
        Page 253
        Page 254
        Page 255
        Page 256
        Page 257
        Page 258
        Page 259
        Page 260
        Page 261
        Page 262
        Page 263
        Page 264
        Page 265
        Page 266
        Page 267
        Page 268
        Page 269
        Page 270
        Page 271
        Page 272
        Page 273
        Page 274
        Page 275
        Page 276
        Page 277
        Page 278
        Page 279
        Page 280
        Page 281
        Page 282
        Page 283
        Page 284
        Page 285
        Page 286
        Page 287
        Page 288
        Page 289
        Page 290
        Page 291
        Page 292
        Page 293
        Page 294
        Page 295
        Page 296
        Page 297
        Page 298
        Page 299
        Page 300
        Page 301
        Page 302
        Page 303
        Page 304
        Page 305
        Page 306
        Page 307
        Page 308
        Page 309
        Page 310
        Page 311
        Page 312
        Page 313
        Page 314
        Page 315
        Page 316
        Page 317
        Page 318
        Page 319
        Page 320
        Page 321
        Page 322
        Page 323
        Page 324
        Page 325
        Page 326
        Page 327
        Page 328
        Page 329
        Page 330
        Page 331
        Page 332
        Page 333
        Page 334
        Page 335
        Page 336
        Page 337
        Page 338
        Page 339
        Page 340
        Page 341
        Page 342
        Page 343
        Page 344
        Page 345
        Page 346
        Page 347
        Page 348
        Page 349
        Page 350
        Page 351
        Page 352
        Page 353
        Page 354
        Page 355
        Page 356
        Page 357
        Page 358
        Page 359
        Page 360
        Page 361
        Page 362
        Page 363
        Page 364
        Page 365
        Page 366
        Page 367
        Page 368
        Page 369
        Page 370
        Page 371
        Page 372
        Page 373
        Page 374
        Page 375
        Page 376
        Page 377
        Page 378
        Page 379
        Page 380
        Page 381
    Illustrations
        Page 382
        Page 383
        Page 384
        Page 385
        Page 386
        Page 387
        Page 388
        Page 389
        Page 390
        Page 391
        Page 392
        Page 393
        Page 394
        Page 395
        Page 396
        Page 397
        Page 398
        Page 399
        Page 400
        Page 401
        Page 402
        Page 403
        Page 404
        Page 405
        Page 406
        Page 407
        Page 408
        Page 409
        Page 410
        Page 411
        Page 412
        Page 413
        Page 414
        Page 415
        Page 416
        Page 417
        Page 418
        Page 419
        Page 420
        Page 421
        Page 422
        Page 423
        Page 424
        Page 425
        Page 426
        Page 427
        Page 428
        Page 429
        Page 430
        Page 431
        Page 432
        Page 433
        Page 434
        Page 435
        Page 436
        Page 437
        Page 438
        Page 439
        Page 440
        Page 441
        Page 442
        Page 443
        Page 444
        Page 445
        Page 446
        Page 447
        Page 448
        Page 449
        Page 450
        Page 451
        Page 452
        Page 453
        Page 454
        Page 455
        Page 456
        Page 457
        Page 458
        Page 459
        Page 460
        Page 461
        Page 462
    Appendix
        Page 463
        Page 464
        Page 465
        Page 466
        Page 467
        Page 468
        Page 469
        Page 470
        Page 471
        Page 472
        Page 473
        Page 474
        Page 475
        Page 476
        Page 477
        Page 478
        Page 479
        Page 480
        Page 481
        Page 482
        Page 483
        Page 484
        Page 485
        Page 486
        Page 487
        Page 488
        Page 489
        Page 490
        Page 491
        Page 492
        Page 493
        Page 494
        Page 495
        Page 496
        Page 497
        Page 498
        Page 499
        Page 500
        Page 501
        Page 502
        Page 503
        Page 504
        Page 505
        Page 506
        Page 507
        Page 508
        Page 509
        Page 510
        Page 511
        Page 512
        Page 513
        Page 514
        Page 515
        Page 516
        Page 517
        Page 518
        Page 519
        Page 520
        Page 521
        Page 522
        Page 523
        Page 524
        Page 525
        Page 526
        Page 527
        Page 528
        Page 529
        Page 530
        Page 531
        Page 532
        Page 533
        Page 534
        Page 535
        Page 536
        Page 537
        Page 538
        Page 539
        Page 540
        Page 541
        Page 542
        Page 543
        Page 544
        Page 545
        Page 546
        Page 547
        Page 548
        Page 549
        Page 550
        Page 551
        Page 552
        Page 553
        Page 554
        Page 555
        Page 556
        Page 557
        Page 558
        Page 559
        Page 560
        Page 561
        Page 562
        Page 563
        Page 564
        Page 565
        Page 566
        Page 567
        Page 568
        Page 569
        Page 570
        Page 571
        Page 572
        Page 573
        Page 574
        Page 575
        Page 576
        Page 577
        Page 578
        Page 579
        Page 580
        Page 581
        Page 582
        Page 583
        Page 584
        Page 585
        Page 586
        Page 587
        Page 588
        Page 589
        Page 590
        Page 591
        Page 592
        Page 593
        Page 594
        Page 595
        Page 596
        Page 597
        Page 598
        Page 599
        Page 600
        Page 601
        Page 602
        Page 603
        Page 604
        Page 605
        Page 606
        Page 607
        Page 608
        Page 609
        Page 610
        Page 611
        Page 612
        Page 613
        Page 614
        Page 615
        Page 616
    Bibliography
        Page 617
        Page 618
        Page 619
        Page 620
        Page 621
        Page 622
        Page 623
        Page 624
        Page 625
        Page 626
        Page 627
        Page 628
        Page 629
        Page 630
        Page 631
        Page 632
        Page 633
        Page 634
        Page 635
        Page 636
        Page 637
        Page 638
        Page 639
        Page 640
        Page 641
        Page 642
        Page 643
        Page 644
        Page 645
        Page 646
        Page 647
        Page 648
        Page 649
        Page 650
        Page 651
        Page 652
        Page 653
        Page 654
        Page 655
        Page 656
        Page 657
        Page 658
    Biographical sketch
        Page 659
        Page 660
        Page 661
    Copyright
        Copyright
Full Text













THE SCARAB BEETLES OF FLORIDA
(COLEOPTERA: SCARABAEIDAE)
PART I. THE LAPAROSTICTI
(SUBFAMILIES: SCARABAEINAE,
APHODIINAE, HYBOSORINAE, OCHODAEINAE,
GEOTRUPINAE, ACANTHOCERINAE)





By
ROBERT EUGENE WOODRUFF


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY









UNIVERSITY OF FLORIDA
December, 1967













PREFACE


This study was begun in 1958 when I joined the staff of the

Division of Plant Industry, Florida Department of Agriculture (formerly

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.

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), "Lepidoptera of Florida" (Kimball,

1965), and "Armored Scales of Florida", (Dekle, 1965). Other volumes are

in preparation as a part of a series on the Arthropods of Florida, being

published by the Division of Plant Industry. This study of the

Scarabaeidae of Florida will be published as a part of that series.

There are those who consider faunal studies as one of the lesser

forms of research, but their viewpoint is probably biased toward some

other specialized study. Herbert Osborne, one of the greatest pioneer

entomologists, once said (1912:63): "While the preparation of such lists

may by some be considered as a rather easy part of entomological investi-

gations, it appears to me that accurately done work of this kind

ii







[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

interest." His statement is equally applicable 30 years later.

The value of such faunal studies is often not appreciated except by

the relatively limited number of persons involved with the organisms.

However, the demand for such a volume often exceeds that for strict

taxonomic revisions. A good example is Blatchley's (1910) monumental

"Coleoptera of Indiana." Even though it is, of necessity, mostly a

compilation and is far out of date, it is one of the most sought after

publications on beetles today.

Florida offers an ideal state for faunal studies because it is a

peninsula and is easily delimited on three sides. It is an important

area from a zoogeographic standpoint because of the proximity of the

West Indies and because 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.

The present study is not taxonomic, nor is it ecological, biological,

ethological or zoogeographical, but it is faunal in nature, encompassing

all of the above aspects. It is a compilation of existing information,

with original data interspersed. It should be primarily useful to those

who wish to identify Florida specimens and then to determine what is


iii








known about that species. If it serves this purpose and stimulates

additional studies, I will be adequately repayed for the effort involved

in its preparation.













ACKNOWLEDGMENTS


As an entomologist with the Division of Plant Industry, Florida

Department of Agriculture, I have been able to pursue several aspects

of this work over a period of ten years. For their encouragement and

understanding I wish to thank the following administrators of this

organization: Doyle E. Conner, Commissioner of Agriculture; H. L. Jones

and the late W. G. Cowperthwaite, Directors of the Division of Plant

Industry; and H. A. Denmark, Chief of the Entomology Section.

I have also had the benefit of close association with my

colleagues of 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.

I am greatly indebted to several museums for the generous loan of

specimens and for the use of their facilities during personal visits.

The 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 abbreviation: Academy of Natural Sciences

of Philadelphia (ANSP), Dr. S. Roback; American Museum of Natural History

(AMNH), Dr. M. A. Cazier, Mrs. P. Vaurie; Canadian National Collection

(CNC), Dr. H. F. Howden; Chicago Natural History Museum (CMNH), Dr. R. L.

Wenzel, Dr. H. S. Dybas; *Florida State Collections of Arthropods (FSCA),

*The Florida State Collections of Arthropods is composed of several collec-
tions 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 collection (REW)
of Scarabaeidae is located with this collection which is housed by the
Division of Plant Industry, Florida Dept. Agr.

V








Dr. H. V. Weems, Jr.; Museum of Comparative Zoology (MCZ), Dr. P. J.

Darlington, Dr. J. F. Lawrence; North Carolina State University (NCS),

Dr. T. B. Mitchell; Ohio State University (OSU), Prof. J. N. Knull,

Mr. F. J. Moore, Dr. C. A. Triplehorn; Purdue University, Blatchley

Collection (PU), Dr. Leland Chandler; United States National Museum (USNM),

Mr. 0. L. Cartwright; University of Florida (UF), Dr. T. J. Walker,

Dr. L. C. Kuitert; University of Miami (UM), Dr. H. F. Strohecker;

University of Michigan Museum of Zoology (UMMZ), Dr. R. D. Alexander, and

Dr. T. H. Hubbell.

I am also indebted to the following individuals who loaned or

donated material from their private collections: Bernard Benesh, L. J.

Bottimer, 0. 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, 0. L. Cartwright,

H. F. Howden, P. 0. Ritcher, and P. Vaurie.

Light taps were one of the greatest sources of material for this

study. Through the aid of the following individuals, samples were

obtained from a network of stations throughout the state: P. E. Briggs,

R. E. Brown, F. A. Buchanan, T. W. Boyd, F. S. Blanton, E. M. Collins, Jr.,

E. E. Crooks, G. W. Desin, C. F. Dowling, H. M. Faircloth, E. H. Frederic,

J. C. Hanlon, E. I. Hazard, L. A. Hetrick, J. Hayward, E. W. Holder, Jr.,

E. G. Kelsheimer, R. L. King, J. H. Knowles, M. Lutrick, E. S. 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, J. E. Porter, W. C. Rhoades,








R. W. Swanson, W. B. Tappan, L. W. Taylor, J. W. Wilson, and D. 0.

Wolfenbarger.

I wish to thank a great many individuals, especially inspectors of

the Division of Plant Industry and U. S. Department of Agriculture, for

providing assistance and specimens. Space does not permit a full listing

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, and E. L. Wells. I

wish to thank Miss A. L. Henley, Librarian of the Division of Plant

Industry for assistance in locating references. I am grateful to Dr. W.

Auffenberg, Curator of Natural Sciences, Florida State Museum, for providing

unpublished data and the distribution map of the gopher tortoise in

Florida.

I wish to thank my supervisory committee for their assistance,

especially for their indulgence with the lengthy manuscript: Drs. 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 for her encouragement

and great care and perseverance in typing the manuscript.














TABLE OF CONTENTS

Page
Preface ........................................................... ii
Acknowledgments........... .............. ....................... v
List of Figures.................................................... xi
Abstract...... ............................................. xvii

Hisntrorical Resduction.................................................... 2
Historical Resume*.**********************************... ....... 2
Collecting Techniques........................................... 4
Format of Presentation................7........................ 7
Florida Biogeography.......................................... 10
General Account of the Family. ....................................... 14
Taxonomy........................................................ 14
Morphology ....................... o................... ........... 18
Biology........... ............................................. 22
Zoogeography................................................. 28
Paleontology...** *******.*.....*............................. .............. 30
Economic Importance....................................... ...... 31
Preliminary Checklist of Florida Scarabaeidae...................... 35
Systematic Account................................................. 44
Key to the Florida Subfamilies of Scarabaeidae.................. 44
Subfamily Scarabaeinae. .................................. ... 48
Key to the Florida Tribes and Genera of Scarabaeinae....... 50
Tribe Scarabaeini......*.....................*...... .... 53
Genus Deltochilum..... ..... ********* *******.......****......... 55
Genus Canthon.......................................... 61
Genus Boreocanthon..................................... 74
Genus Melanocanthon.................................... 79
Genus Glaphyrocanthon........ ................ ......*********************.. 85
Genus Pseudocanthon.............................*.... 89
Tribe Coprini............................................ 93
Genus Ateuchus*.**************************************. 95
Genus Dichotomius...................................... 101
Genus Copris........................................... 106
Genus Phanaeus.............*...............* ........... 118
Tribe Onthophagini....................................... 126
Genus Onthophagus ..................................... 126
Tribe Oniticellini....................................... 157
Genus Oniticellus.................................... 159
Subfamily Aphodiinae......... .....*......................... 164
Key to the Florida Genera of Aphodiinae.................... 167
Genus Aphodius....*....*..*............................ 170
Genus Myrmecaphodius................................... 222
Genus Euparia.......................................... 225
Genus Ataenius........................................ 228
Genus Pseudataenius.................................... 273


viii








Page
Genus Aphotaenius ..... ................................. 275
Genus Psammodius....................................... 279
Genus Pleurophorus ... ................................. 288
Genus Rhyssemus........................................ 294
Subfamily Hybosorinae........................................ 299
Genus Hybosorus........................................ 300
Subfamily Ochodaeinae...................................... 304
Genus Ochodaeus........................................ 306
Subfamily Geotrupinae................ .................... 316
Key to the Florida Tribes and Genera of Geotrupinae........ 319
Tribe Bolboceratini .......... ... .......... ........... 320
Genus Bolboceras....................................... 321
Genus Bradycinetulus.......... ...................... 324
Genus Bolbocerosoma................................... 327
Genus Eucanthus.............. ......................... 330
Tribe Geotrupini........... ............................ 337
Genus Geotrupes........................................ 338
Genus Peltotrupes...................................... 346
Genus Mycotrupes....................................... 355
Subfamily Acanthocerinae................................. 369
Key to the Florida Genera and species of Acanthocerinae.... 371
Genus Acanthocerus..................................... 372
Genus Cloeotus......................................... 374
Illustrations (see List of Figures p. xi).......................... 382
Appendices...................................... ................... 463
1. Deltochilum gibbosum gibbosum (Fabricius)................... 465
2. Canton pilularius (Linnaeus) .............................. 466
3. Canthon vigilans LeConte.................................... 468
4. Boreocanthon depressipennis (LeConte)..................... 470
5. Boreocanthon probus (Germar)............o................. 472
6. Melanocanthon bispinatus (Robinson)........................ 473
7. Melanocanthon granulifer (Schmidt)........................ 474
8. Melanocanthon punctaticollis (Schaeffer)................... 475
9. Glaphyrocanthon viridis (Beauvois)......................... 476
10. Pseudocanthon perplexus (LeConte).......................... 477
11. Ateuchus lecontei (Harold).................................. 478
12. Dichotomius carolinus (Linnaeus)............................ 483
13. Copris inemarginatus Blatchley................. 485
14. Copris minutes (Drury).................................. 486
15. Phanaeus igneus floridanus d'lsouieff................... 490
16 FPhanaeus vindex MacLeay ................................ 494
17. Onthophagus hecate blatchleyi Brown......................... 497
18. Onthophagus oklahomensis Brown.............................. 500
19. Onthophagus pennsylvanicus Harold........................... 502
20. Onthophagus striatulus floridanus Blatchley.................. 503
21. Onthophagus tuberculifrons Harold........................... 505
22. Aphodius aegrotus Horn....................................... 510
23. Aphodius campestris Blatchley............................... 512
24. Aphodius crassulus Horn.........................*........... 518
25. Aphodius cuniculus Chevrolat....*************. *****........ 519
26. Aphodius fimetarius (Linnaeus)..................... ....... 522
27. Aphodius floridanus Robinson................................ 523
28. Aphodius laevigatus Haldeman.............. ............... 524








Page
29. Aphodius lividus (Olivier)............... ................ 525
30. Aphodius parcus Horn...... ............................. 532
31. Aphodius rubeolus Beauvois................................. 535
32. Aphodius stupidus Horn.................................... 537
33. Aphodius troglodytes Hubbard............................ 538
34. Euparia castanea Serville ............................ 539
35. Myrmecaphodius proseni Martinez ........................... 543
36. Ataenius alternatus (Melsheimer)......................... 544
37. Ataenius cylindrus Horn.................................... 550
38. Ataenius erratus Fall.................................... 554
39. Ataenius exiguus Brown................................... 555
40. Ataenius fattigi Cartwright............................. 558
41. Ataenius gracilis (Melsheimer)........................... 561
42. Ataenius imbricatus (Melsheimer).......*................... 566
43. Ataenius miamii Cartwright..... ................... .... 569
44. Ataenius picinus Harold.................................... 570
45. Ataenius platensis Blanchard........... ............... 571
46. Ataenius rudellus Fall......................... .... ... 57
47. Ataenius saramari Cartwright............................... 575
48. Ataenius simulator Harold................................. 576
49. Ataenius spretulus (Haldeman)...............* .............. 583
50. Ataenius strigicauda Bates............................... 586
51. Psammodius malkini Cartwright.............................. 587
52. Pleurophorus longulus Cartwright ........................... 590
53. Hybosorus illigeri Reiche.................................. 594
54. Ochodaeus frontalis LeConte............................... 600
55. Bolbocerus floridensis (Wallis) .......................... 603
56. Bradycinetulus ferrugineus (Beauvois)..................... 604
57. Bolbocerosoma hamatum Brown............ .................. 605
58* Eucanthus impressus Howden .............................. 606
59. Eucanthus subtropicus Howden.......................... 607
60. Geotrupes egeriei Germar.................................. 610
61. Peltotrupes profundus Howden.......... ........ ......... 612
62. Mycotrupes gaigei Olson & Hubbell.......................... 614
63. Cloeotus aphodioides (Illiger)............................. 615
64. Cloeotus globosus (Say).................................... 616
Bibliography. ............................................. ..... 617
Biographical Sketch...................................... ....... 659














LIST OF FIGURES

1 32. Habitus drawings of one member of each genus of Florida
Laparosticti.
Page
1. Phanaeus igneus floridanus d'Olsoufieff.................... 383
2. Deltochilum g. gibbosum (Fab.)............................ 384
3. Canthon pilularius (L........... ............... 384
4. Boreocanthon depressipennis (Lec.)......................... 384
5. Melanocanthon punctaticollis (Schffr.)..................... 384
6. Glaphyrocanthon v. viridis.............................. 385
7. Pseudocanthon perplexus (Lec.)............................. 385
8. Ateuchus lecontei Ha.)................................... 385
9. Dichotomius carolinus (L.) ........ ................. o...... 385
10. Copris minutes (Drury).................................... 386
11. Onthophagus p. polyphemi Hub. ............................. 386
12. Oniticellus cubiensis Lap. ................................ 386
13. Aphodius fimetarius (L.)................................... 386
14. Myrmecaphodius proseni Mart. ...............*.....**....... 387
15. Euparia castanea Serv. .................................... 387
16. Ataenius saramari Cartwr. ............*.................... 387
17. Aphotaenius carolinus (Van D.)......................... 387
18. Pseudataenius socialis (Horn)............................. 388
19. Rhyssemus scaber Hald. ................................... 388
20. Psammodius malkini Cartwr. ............................... 388
21. Pleurophorus longulus Cartwr ............................. 388
22. Hybosorus illigeri Reiche............................... 389
23. Ochodaeus frontalis Lee. ....................... ....... 389
24. Bolboceras floridensis (Wallis)............................ 389
25. Bradycinetulus ferrugineus (Beauv.)...................... 389
26. Bolbocerosoma hamatum Brown................................ 390
27. Eucanthus subtropicus Howden.............................. 390
28. Geotrupes egeriei Germar........................ ......... 390
29. Peltotrupes profundus Howden............................. 390
30. Mycotrupes gaigei Ols. & Hub............................. 391
31. Cloeotus globosus (Say).................................... 391
32. Acanthocerus aeneus MacL. ................................. 391

33 116. Distribution maps of Florida Laparosticti.

33. Deltochilum g. gibbosum (Fab.)............................. 392
34. Canthon chalcites Hald.)................................ 392
35. Canthon pilularius (L.)................................ 393
36. Canthon vigilans Lec. ................... ................ 393
37. Boreocanthon depressipennis (Lec.)......................... 394







Page
38. Boreocanthon probus (Germar)............. ... .... 394
39. Melanocanthon bispinatus (Rob.)......... ..............** 395
40. Melanocanthon granulifer (Schm.) ....*.. 395
41. Melanocanthon punctaticollis (Schffr.).....................396
42. Glaphyrocanthon v. viridis (Beauv.)............... **o.. 396
43. Pseudocanthon perplexus (Lec.).......*....... .. ,,, ., 397
4A. Ateuchus lecontei (Har.) .................................. 397
45. Dichotomius carolinus (L.)............................... 398
46. Copris gopheri Hubbard............................. ... 398
47. Copris howdeni Matth. & Halfft., inemarginatus Blatch. ..... 399
48. Copris minutus (Drury)................. .,,,.......... 399
49. Phanaeus i. igneus MacL., i. floridanus d'Ols.............. 400
50. Phanaeus index MacL. ***................*...... ....... 400
51. Onthophagus aciculatulus Blatch., concinnus Lap............ 401
52. Onthophagus depressus Har. *********.................... .... 401
53. Onthophagus hecate blatchleyi Brown..........**... .... 402
54. Onthophagus oklahomensis Brown.....*..**......... ........... 402
55. Onthophagus o. orpheus (Panz.).............................. 03
56. Onthophagus pennsylvanicus Har.................... ... 403
57. Onthophagus p. polyphemi Hub., p. sparsisetosa Howd. &
Cartwr .. ........................................404
58. Onthophagus striatulus floridanus Blatch. ................. 404
59. Onthophagus subaeneus (Beauv.).******.......................... 405
60. Onthophagus tuberculifrons Har............................. 405
61. Aphodius aegrotus Horn............**.....* ........, 406
62. Aphodius bicolor Say........*...........................*... 406
63. Aphodius campestris Blatch. ......................... .... 407
64. Aphodius crassulus Horn..oo ********......................... 407
65. Aphodius cuniculus Chevr. ..........t....................... 408
66. Aphodius fimetarius (L.)................ 408
67. Aphodius floridanus Rob. .........** ....................... 409
68. Aphodius haemorrhoidalis (L.)................*............. 409
69. Aphodius laevigatus Hald ................**.............. 410
70. Aphodius lividus (Oliv.).................... ............ 410
71. Aphodius lutulentus Hald. ....**............................ 411
72. Aphodius parcus Horn.**.....*********... .................... 41l
73. Aphodius rubeolus (Beauv.)..*............................... 412
74. Aphodius stupidus Horn.................................... 412
75. Aphodius troglodytes Hubbard...........,.4.................. 13
76. nyrmecaphodius proseni Mart. ......... ....................... 4
77. Euparia castanea Serv. *************.... **................. 414
78. Ataenius alternates (Melsh.)... .......*... ............... 414
79. Ataenius brevinotus Chapin........... .................... 415
80. Ataenius cylindrus Horn........................ .... ........ 415
81. Ataenius erratus Fall........................... ........ 416
82. Ataenius exiguus Brown...********************** ........... 416
83. Ataenius fattigi Cartwr. .. ****....**.......................... 417
84. Ataenius gracilis (Melsh.).................................. 417
85. Ataenius imbricatus (Melsh.).....*.......................... 418
86. Ataenius insculptus Horn.............***..................... 418
87. Ataenius linelli Cartwr., luteomargo Chapin, ovatulus Horn.. 419
88. Ataenius miamii Cartwr. *****************.....................* 419
89. Ataenius picinus Har. *******************................... 420


xii








Page
90. Ataenius platensis Blanch. ................................. 420
91. Ataonius rudellus Fall.................................... 421
92. Ataenius saramari Cartwr. ............................... 421
93. Ataenius simulator Har. ....................... ........ 422
94. Ataenius spretulus Hald. .....*.........*................. 422
95. Ataenius strigicauda Bates.................................. 423
96. Pseudataenius socialis (Horn) .............................. 423
97. Psammodius armaticeps Fall................................. 424
98. Psammodius bidens Horn.............. .................. ... 424
99. Psammodius cruentus Har. ..........................*........ 425
100. Psammodius malkini Cartwr. ................... ............. 425
101. Pleurophorus longulus Cartwr. .....*.................. .. ... 426
102. Hybosorus illigeri Reiche..............*.................... 426
103. Ochodaeus frontalis Lec. ................* ............... 427
104. Bolboceras floridensis (Wallis)............................. 427
105. Bradycinetulus ferrugineus (Beauv.).*....................... 428
106. Bolbocerosoma hamatum Brown. ............................. 428
107* Eucanthus alutaceus Cartwr. ................................ 429
108. Eucanthus impressus Howd. .......* ......................... 429
109. Eucanthus subtropicus Howd. .............*............... 430
110. Geotrupes b. blackburnii (Fab.)............................. 430
111. Geotrupes egeriei Germar.............*...................... 431
112. Mycotrupes gaigei 0. & H., cartwright 0. & H., pedester
Howd. ...................... 431
113. Peltotrupes profundus Howd., and youngi Howd. .............. 432
114. Cloeotus aphodioides (II.)..... 432
115. Cloeotus globosus (Say).. ............................ 433
116. Acanthocerus aeneus MacL. .................................. 433

117. Distribution map of the gopher tortoise (Gopherus
polyphemus (Daudin))......... .... *o*.*...*..........*.. 434
118. Precipitation in Florida......................*............. 434

119 138. Photographs of habitats of Florida Laparosticti.

119. Dune habitat at St. Andrews State Park, Fla. ............... 435
120. Close up of a rosemary bush (Ceratiola ericoides)........... 435
121. Habitat of Mycotrupes cartwrighti Ols. & Hub. .............. 436
122. Mounds of the pocket gopher............................... 436
123. Bait traps in place.... .................................. 437
124. Geotrapes egeriei Germn emerging from pupal cell............ 437
125. Habitat at Cape Sable, Fla. ................................ 438
126. Beach habitat at Cape Sable, Fla. .......................... 438
127. Packrat habitat at Florida Caverns State Park............... 439
128. Packrat droppings .......................... * *.***.*.***.. 439
129. Stick nest of Packrats at Key Largo, Fla. .................. 440
130. Packrat nest after removal of stick covering................ 440
131. View from bluffs at Torreya State Park, Fla. ............... 441
132. "Push-up" of Peltotrupes profundus Howden................... 441
133. Dichotomius carolinus (L.) larva repairing cell............. 442
134. Dichotomius carolinus (L.) larva, showing "hump-backed"
appearance..................................................******** 442


xiii







Page
135. Pasture containing imported fire ant mounds................ 443
136. Cross section of an imported fire ant nest............ ,.... 443
137. Burrowing owl at the entrance to its burrow................ 444
138. Burrowing owl pellets containing insect remains............ 444

139 204. Outline drawings of morphological characters.

139. Dorsal view of head of Ochodaeus frontalis Lec. .......... 446
140. Dorsal view of head of Ochodaeus musculus Say.............. 446
141. Pectinate inner spur on left middle tibia of male
Ochodaeus frontalis Lec. ...................... ........*. 446
142. Pectinate inner spur on left middle tibia of male Ocho-
daeus frontalis Lee. ............................... 446
143. Pectinate inner spur on left middle tibia of female
Ochodaeus frontalis Lec. ...............................*** 446
144. Caudal view of pygidium and propygidium of Ochodaeus
frontalis Lec. ................ .........*****............******** 446
145. Ventral view of the right posterior femur of a male
Ochodaeus frontalis Lec. ............. ...... .*********** 446

146 153. Hybosorus illigeri Reiche.

146. Left lateral view of male genitalia.................................... 448
147. Right lateral view of male genitalia....................... 448
148. Dorsal view of male genitalia.............................. 448
149. Ventral view of left posterior leg4........................ 448
150. Dorsal view of left antenna................................ 448
151. Caudal view of antennal club............................... 448
152. Dorsal view of head...448...................... 448
153. Dorsal view of left anterior tibia and tarsus.............. 448

154 166. Acanthocerinae.

154. Dorsal view of head of Acanthocerus aeneus MacL............ 450
155. Dorsal view of left elytral margin of Cloeotus
aphodioides (Ill.)......................................... 450
156. Dorsal view of head of Cloeotus globosus (Say)....... ..... 450
157. Dorsal view of head of Cloeotus aphodioides (Ill.)......... 450
158. Dorsal view of left elytral margin of Cloeotus
globosus (Say)............................................ 450
159. Dorsal view of left anterior pronotal angle of
Acanthocerus aeneus MacL. ................................. 450
160. Dorsal view of left anterior pronotal angle of Cloeotus
globosus (Say)..... .............. **********************... 450
161. Ventral view of right antenna of Cloeotus aphodioides
(Ill.)................................................... 452
162. Ventral view of left anterior tibia and tarsus of C.
aphodioides (Ill.).....................*...*................ 452
163. Ventral view of left posterior femur and tibia of
Acanthocerus aeneus MacL. ................................. 452
164. Cross section of posterior tibia of A. aeneus MacL. ....... 452
165. Ventral view of left posterior femur, tibia, and tarsus
of Cloeotus aphodioides (Ill.).............************** 452
166. Cross section of posterior tibia of C. aphodioides (nIl.).. 452


xiv








167 174. Larva of Cloeotus aphodioides (Ill.). Page

167. Frontal view of head...................... ................ 454
168. Ventral view of epipharynx................................. 454
169. Dorsal view of left maxilla................................ 454
170. Dorsal view of left mandible.............................. 454
171. Dorsal view of right mandible.............................. 454
172. Venter of last abdominal segment........................... 454
173. Portion of the stridulatory area on the metathoracic leg... 454
174. Thoracic spiracle.......................................... 454

175. Dorsal view of the abdomen of Euphoria sepulchralis
(Fab.)..... .... ......... ....... ......................... 456
176. Dorsal view of the abdomen of Copris minutus (Drury)....... 456
177. Lateral view of the abdomen of Euphoria sepulchralis
(Fab.).. ......... ................ ....................... 456
178. Ventral view of the abdomen of Cloeotus aphodioides
(Ill.)..................................................... 456
179. Ventral view of the abdomen of Geotrupes egeriei Germ. .... 456
180. Dorsal view of right posterior leg of Aphodius
fimetarius (L.)........4........................... ...... 456
181. Dorsal view of the right posterior leg of Ataenius
alternatus (Melsh.).........***...................... 456
182. Lateral body outline of Cloeotus globosus (Say)............ 458
183. Lateral body outline of Ataenius alternates (Melsh.)....... 458
184. Dorsal view of the right antenna of a male Phanaeus
igneus floridanus d'Ols* *******************************............... 458
185. Dorsal view of the right antenna of a male Polyphylla
pubescens Cartrw. ................. ....................... 458
186. Ventral view of the right posterior tarsus of Phanaeus
igneus floridanus d'Ols. .................................. 458
187. Lateral view of the tip of the left posterior tarsus of
Phyllophaga latifrons (Lee.) ... o......* *... ................ 458
188. Dorsal view of the right anterior tibia of a male
Phanaeus igneus floridanus dOls. ************************* 458

189 195. Canthon.

189. Dorsal view of the head of Canthon vigilans Lec. .......... 460
190. Dorsal view of the head of Canthon pilularius (L.)......... 460
191. Lateral view of the male genitalia of Canthon vigilans
Lee. ................................... ........4........ 460
192. Lateral view of the male genitalia of Canthon pilularius
(L.).................................................... 460
193. Anterior right tibial spur of male Canthon chalcites
(Hald.)....... ...................................... 460
194. Anterior right tibial spur of male Canthon chalcites
(Hald.)....................................... 460
195. Anterior right tibial spur of female Canthon chalcites
(Hald.)...............*.................................. 460

196. Dorsal view of right posterior tibia of Dichotomius
carolinus (L.)**********************************........... 462







Page
197. Dorsal view of right posterior tibia of a male
Deltochilum gibbosum (Fab.)............................... 462
198. Dorsal view of anterior tibia of a female Deltochilum
gibbosum (Fab.).......................................... 462
199. Dorsal view of right antenna of Geotrupes egeriei Gem. .. 462
200. Dorsal view of right antenna of Bolboceras floridensis
(Wallis).................................................. 462
201. Dorsal view of right posterior tibial apex and tarsus of
Pleurophorus longulus Cartwr. .... ...................... 462
202. Dorsal view of right posterior tibial apex and tarsus of
Psamnodius malkini Cartwr. ............................... 462
203. Dorsal view of right posterior tibia and tarsus of
Melanocanthon granulifer Schmidt.......................... 462
204. Dorsal view of right posterior tibia and tarsus of
Boreocanthon depressipennis (Lec.)........................ 462









Abstract of Dissertation Presented to the Graduate Council in
Partial Fulfillment of the Requirements for the Degree of
Doctor of Philosophy


THE SCARAB BEETLES OF FLORIDA (COLEOPTERA: SCARABAEIDAE)
PART I. THE LAPAROSTICTI
(SUBFAMILIES: SCARABAEINAE, APHODIINAE, HYBOSORINAE,
OCHODAEINAE, GEOTRUPINAE, ACANTHOCERINAE)


By

Robert Eugene Woodruff

December, 1967


Chairman: L. A. Hetrick
Major Department: Entomology


In this faunal study 240 species and subspecies of Scarabaeidae

&r'e recorded from Florida. Only the Laparosticti, representing 106

spocies and subspecies, are treated in detail, although a checklist is

provided for the entire family. Keys are presented for the identi-

fication of all taxonomic units, with many of the morphological

characters illustrated by line drawings. 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 over 78,638 specimens are presented in tabular

form in the appendices. The bibliography contains 585 references.

Twenty habitat photographs are included.


xvii









Three species are recorded for the first time in the U. S.:

Myrmecaphodius proseni Martinez, previously known from Argentina;

Ataenius brevinotus Chapin, previously known from Cuba; and Ataenius

luteomargo Chapin, known from several islands in the West Indies. 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

vindex cyanellus Robinson and Phanaeus difformis magnificens Robinson

are synonymized under Phanaeus vindex MacLeay.


xviii













INTRODUCTION


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).

The taxonomy of the U. S. species is relatively stable-that is,

few species are described each year. Several taxonomists (e.g. Cartwright,

Howden, Matthews, Vaurie, and myself) are actively engaged in revising

certain portions of the family. Numerous generic revisions, during the

past 15 years, have clarified many of the problems, but much remains to

be done.

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 needed." Obviously such an undertaking

would be a worthwhile but extremely time consuming proposition. Since

it appears that such an opus is not foreseen in the near future, the

next best approach would seem to be limited faunal studies.

Many of the gaps in our knowledge become especially apparent

after a detailed local faunal study such as this one. Because of the

large number of species involved, it has not been possible to pursue






2

and clarify many of the problems encountered. However, I have tried to

point them out in the species discussions, in the hope that they will

stimulate future work along these lines.

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 can be divided into two groups-the

Laparosticti and the Pleurosticti. The former is characterized by having

the abdominal spiracles situated in a line on the membrane between the

sternites and tergites, and includes the dung feeding and scavenger species.

These are represented in Florida by the following six subfamilies:

Scarabaeinae, Aphodiinae, Geotrupinae, Hybosorinae, Ochodaeinae, and

Acanthocerinae. The Pleurosticti are characterized by having most of the

abdominal spiracles situated on the upper portion of the sternites, and

include primarily the plant feeders and chafers. These are represented

in Florida by the following four subfamilies: Melolonthinae, Rutelinae,

Cetoniinae, and Dynastinae.

Due to limitations of time and space this dissertation will deal

only with the Laparosticti. However, data have been accumulated 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.


Historical Resume


There have been scattered records of Florida Scarabaeidae 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 Suwanee

Springs. Hamilton (1894) recorded 11 species from Lake Worth and added

six 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. Wickham (1909) reported 26 species from seven Florida

localities.

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 undescribed (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) recorded 13 species

of Serica. Frost (1964) recorded 23 species from blacklight traps at

Archbold Biological Station (Highlands Co.). In the present study I

have recorded 240 species and subspecies, excluding all previous erroneous

records, but including 11 questionable species (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: Blatchley (1910) in








his "Coleoptera of Indiana"; Dawson (1922), the "Scarabaeidae of

Nebraska"; 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.


Collecting Techniques


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 techniques 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 sometimes difficult to locate dung of such wild animals,

and most of the beetles associated with these hosts remain rare in

collections. Although some species complete 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, consisted of any convenient

container (usually tin cans or pint glass jars) sunk into the ground,

level with the top (Fig. 123). The soil was packed tightly around the

rim to enable the beetles to crawl in. In the case of liquid baits, the

trap was about half filled with the 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 transferred to 70% isopropyl

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 collecting

large numbers of many burrowing groups for which the food habits are un-

known (e.g. Mycotrupes, Peltotrupes). The use of specific attractants

offers a fertile field for future investigation.

Many of the Scarabaeidae are attracted to light, and various

designs of light traps are effective in collecting specimens. Blacklight

(actually ultra violet) is much more effective than white light. The








traps which I have utilized are modifications of those originally de-

signed by the USDA for European chafer (Amphimallon majalis Raz.).

The specimens were collected in 70% isopropyl alcohol placed in the

collecting container. For a discussion of the nature of light sources,

trap designs, and additional information on light trapping, the reader

is referred to the following papers: USDA (1961); Hollingsworth, Hartsock,

and Stanley (1963); Frost (1952, 1958, 1963, 1964, 1966). The advent of

the blacklight has nearly revolutionized 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

abundant specimens for morphological study.

Other mass collecting methods used were modified Berlese or Tulgren

funnels and Malaise traps. Some of the larger species (e.g. Deltochilum,

Dichotomius) have been trapped occasionally in mist nets used for birds

and bats.

Specimens were often floated out of cow dung, leaf litter, plant

roots, etc., by placing the material in a pail of water. Digging was

often the only 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. 132), and they can be followed by in-

serting a straw into the hole and digging along side. Fortunately, these

deep burrows are usually vertical.

Several species, especially Aphodiinae, occur in leaf mold or pine

needles and were found by scraping 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 specimens were found

by pulling up plants in such habitats. They were sometimes found under

boards and the windows of debris behind the high tide marks.

Nearly all specimens were collected in 70% isopropyl alcohol which

permitted easy genitalic dissections. Specimens .can be preserved indef-

initely in this solution and are available for future mounting or

dissection. The immature stages often were found in dung or in the soil

and were usually easily reared in salve tins. For preservation, speci-

mens were killed in boiling water and preserved in 70% isopropyl alcohol.



Format of Presentation


Keys are presented for all of the taxonomic units 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 parenthesis indicating the

rubric to consult for retracing one's steps. The keys are artificial in

the sense that they use characters which are easily seen or by which

specimens are easily separated, without regard to any phylogenetic

scheme. Free use has been made of existing keys, but they have been mod-

ified for Florida forms and will not necessarily apply to other regions.

Many of the keys are original, and every effort has been made to make

them as clear and easy to use as possible. They are all dichotomous,

with the couplets reading parallel. References are made to illustrations

wherever pertinent.






8

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 confused with this term in genetics); by whom and how designated

(e.g. monotypy, subsequent designation, 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 subgenera; status of the

classification (e.g. recently revised, in need of revision, etc.);

variation; subspecies. 6) Distribution and Zoogeography: general

zoogeographic region occupied (e.g. Neotropical, Nearctic, 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,

vertebrate 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: include author, date, and page citation to the

most important papers on the genus; a complete citation is 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







9

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. During the course of preparing the syn-

onymy, a complete list of references was prepared, but it was not included

in the manuscript 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 species, even though they are not found in Florida. Reference

is made to drawings where pertinent. 3) Taxonomic Notes: the status of

the species and reasons for any synonymy are explained. Variation, sub-

speciation, 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. Included 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 records 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 specimens which I have seen are listed

either in the section under "specimens examined" or in the appendices.

The generalized distribution is also based on literature records and on

specimens I have examined. In many cases the records are not sufficient

to present an accurate map, but it was felt that such a summary would

form a basis for future detailed studies. Inadequate as they must be,

these maps are the only ones published for a great number of the species.

5) Biology: this is essentially a "catch 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 and the number of Florida localities

represented. If less than 10 localities are known, the label data is

included here, but if more than 10 localities are involved these data

are presented in the appendices (for which a reference is made). Abbrev-

iations for the collections 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. Only those papers which contain significant data are in-

cluded, and catalogues, or state lists are excluded.

New species which have been discovered are listed here by number

only, with a brief notation about them, pending formal description in

a scientific journal. In cases where these species are to be described

by other workers, a statement is made to this effect. They are not

included in the keys at present, but they will be incorporated before

publication. They are listed at the end of each genus, after the alpha-

betical arrangement of described species.



Florida Biogeography


Studies on a diversity of plant and animal groups 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.







11
Not only was the peninsula a refugium for certain species during glacial

periods, but its unique geographical position and geological history

appear to have been conducive to the establishment of groups of endemic,

closely allied, allopatric species of apparently recent origin (Hubbell,

1961).

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 shorelines. 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 terraces is not firmly established, but several of those

previously considered of Pleistocene age are probably Pliocene or Late

Miocene (Alt and Brooks, 1965)'.

It is also apparent that our present knowledge about the distri-

bution of most insects, including the Scarabaeidae, is too fragmentary

to draw many significant conclusions. In most cases our present

distributional patterns are based on such meagre data that they cannot

be interpreted with any degree of confidence. This situation became very

apparent when I tried to analyze the data on the genera yvcotrupes and

Peltotrupes (see the discussions under these genera). With a few rare

exceptions, it appears too premature 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

incomplete 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 intensively than

any other state, there are large areas which have not been thoroughly

sampled. This is especially 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 patterns can be used as if they

reflected a natural situation.

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 responsible. However,

the data are not sufficient to allow for many generalizations.

Space does not permit a detailed discussion of this interesting,

but complicated, subject. Considerable work being conducted on other

groups of animals and in paleontology will undoubtedly produce new and

interesting data on the past and present distributions of Florida animals.

For additional 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 C-oin (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); Hubbell (1932, 1954, 1961);
Hubbell, Laessle, and Dickinson (1956); Kurz (1942); Laessle (1942,

1958); Landsberg (1949); MacNeill (1951); McCrone (1963); Mitchell





13
(1963); Mount (1963, 1965); Neill (1957); Safford (1919); Schuchert
(1910, 1929, 1935); Schwarz (1888); Vaughan (1910); West and Arnold
(1956); Young (1954).













GENERAL ACCOUNT OF THE FAMILY


Taxonomy

Although the family is one of the better known of the larger

families of beetles, many genera need revising. For example, the large

genera Aphodius and Ataenius have not been revised, even for the U. S.,

since the time of Horn (1887). 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, subtribes, 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 Matthews, 1966).

Many European authors, including Crowson (1955), have treated as

families several of the groups recognized here as subfamilies. Basically

I have followed the arrangement of families and subfamilies of the

superfamily Scarabaeioidea listed by Arnett (1962), with the exception

of recognizing the Trogidae as a full family.








Families Subfamilies of Scarabaeidae

Lucanidae Scarabaeinae
Passalidae Aphodiinae
Scarabaoidae Ochodaeinae
Trogidae Hybosorinae
Geotrupinae
Pleocominae
Glaphyrinae
Acanthocerinae
Melolonthinae
Rutelinae
Dynastinae
Cetoniinae
The arrangement of the superfamily in several major references

is presented in the following discussion. 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 Scarabaeioidea is

arranged as follows:

Families Subfamilies of Scarabaeidae

Lucanidae Coprinae
Passalidae Aegialiinae
Scarabaeidae Aphodiinae
Ochodaeinae
Hybosorinae
Geotrupinae
Pleo cominae
Glaphyrinae
Acanthocerinae
Troginae
Melolonthinae
Rutelinae
Dynastinae
Cetoniinae

Boving and Craighead (1931), in their synopsis of the larval forms of

Coleoptera, treated the superfamily Scarabaeoidea as follows:

Families Subfamilies of Scarabaeidae

Lucanidae Coprinae
Passalidae Aphodiinae
Geotrupidae Glaphyrinae
Trogidae Pleocominae








Subfamilies of Scarabaeidae


Acanthoceridae
Scarabaeidae


Melolonthinae
Sericinae
Macrodactylinae
Rutelinae
Dynastinae
Trichiinae
Valginae
Cetoniinae


Edwards (1949a), in treating the beetles east of the Great Plains,

arranged the Scarabaeoidea as follows:


Families


Subfamilies of Scarabaeidae


Lucanidae
Passalidae
Geotrupidae
Acanthoceridae
Trogidae
Scarabaeidae


Coprinae
Aegialiinae
Aphodiinae
Ochodaeinae
Melolonthinae
Rutelinae
Cetoniinae
Dynastinae


In the "Coleopterorum Catalogus" (1910-1937), a world catalogue authored

by various specialists, the Scarabaeoidea are arranged as follows:


Families


Subfamilies of Scarabaeidae


Lucanidae
Passalidae
Scarabaeidae


Aphodiinae
Aegialiinae
Chironinae
Coprinae
Termitotroginae
Pachypodinae
Pleocominae
Aclopinae
Glaphyrinae
Ochodaeinae
Orphninae
Idiostominae
Hybosorinae
Dynamophinae
Acanthocerinae
Troginae
Melolonthinae
Taurocerastinae
Geotrupinae
Euchirinae
Phaenomerinae
Rutelinae


Families








Families Subfamilies of Scarabaeidae

Dynastinae
Trichiinae
Valginae
Cetoniinae

Crowson (1955), in his treatise on the natural classification of the

families of Coleoptera, treated the Scarabaeoidea as follows:

Families Subfamilies of Scarabaeidae

Lucanidae Taurocerastinae
Passalidae Hybosorinae
Trogidao Orphninae
Acanthoceridae Ochodaeinae
Geotrupidae Allidiostominae
Scarabaeidae s Aphodiinae
Scarabaeinae
Glaphyrinae
Rutelinae
Dynastinae
Cetoniinae
Pachypodinae
Melolonthinae

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

thorough, arrangement is provisionally followed. Where obvious incon-

sistencies have been noted they are pointed out.

Generic names contribute to greater confusion since they are a

part of the binomial nomenclature. Genera do not exist in nature

(although species do) and are merely subjective categories that assist

in organizing the species into groupings. In some cases there are clear-

cut dividing lines between such groups, but more often there is a

gradual transition. Because of the subjective nature of the generic con-

cept, many species have been moved back and forth from one genus to the

next, creating numerous nomeclatural problems, especially homonymy.

This is one reason which led Michener (1964) to suggest the idea of

"uninomial nomenclature."







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 sub-

tribe 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 unavail-

ability of type specimens or literature. Certain others will require

submission 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.



Morphology


Like nearly all other aspects of the Scarabaeidae, the morphology

is quite diverse and difficult to generalize. The family is divided into

two sections-the Laparosticti and Pleurosticti-based on the position

of the abdominal spiracles, the relative positioning of the posterior

legs, as well as differences in biology (i.e. primarily dung feeders and

primarily plant feeders).

The family varies tremendously in size of individuals, from 1.9 mm

(.075 inch) for Pleurophorus longulus Cartwr. to nearly 15 cm






19
(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 a large percentage

of the species are dull colored, the family contains many with beautiful

metallic colors, rivaling the showiness of the most colorful Lepidoptera.

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 Figs. 1-32.

The family has been a popular one with collectors because of their

bright colors and also because of 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 signif-

icance of these structures.

All species have a lamellate antenna club arranged so that the

lamellae can be expanded or appressed 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. 184, 199, 200), but contains as many as seven in the

genus Polyphylla (Fig. 185). The club is variously shaped, from ball-

like in Bolboceras (Fig. 200) to asymmetrical in Phanaeus (Fig. 184) or

somewhat cup-shaped in Hybosorus (Fig. 150). The antennae are the

principal olfactory receptors, 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). 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 beyond the middle of the body, usually nearer the tip of the

abdomen than to the middle pair of legs. As a consequence, the meta-

thorax 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.

The anterior coxae are large, transverse, and sometimes 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 variable in form; they are narrow, curved, and fitted for ball

rolling in the Scarabaeini; flattened and sickle-shaped, and the surface

covered with incised lines in the Acanthocerinae (Fig. 163, 165); 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. 186). Although there is

usually a bisetose onychium, it is wanting in the Acanthocerinae,

Aphodiinae, and some Scarabaeinae.

The mouthparts are highly variable in form, from the nearly mem-

branous type of the Scarabaeinae and Aphodiinae, to the hard, well-

developed mandibles of the Geotrupinae and Dynastinae. The mouthparts

are described in detail by Hardenberg (1907) from examples throughout

the family. Landin (1961) discussed these organs in the Aphodiinae, and






21
Miller (1961) and Halffter and Matthews (1966) considered these structures
in several Scarabaeinae. The maxillary palpi are slender, four-

segmented, with the apical segment the largest. They often contain odor

and, probably, taste receptors. The labial palpi are three-segmented.

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 bacteria."

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), These authors discuss the morphological adaptations for

coprophagy in the adults and larvae of the Scarabaeinae. Becton (1930)

described the alimentary tract of Phanaeus vindex MacL., and Miller

(1961) described 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 Scarabaeinae 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 Coleoptera showing such a

degree of ovarian reduction.

The male genitalia are useful for distinguishing species of certain

genera (e.g. Melanocanthon, Bolbocerosoma, and Phyllophaga). However,

there has been no extensive comparative study within the family. There

are a few studies comparing these organs between families of Coleoptera

(Sharp and Muir, 1912; Snodgrass, 1957).

Although the female genitalia probably are diagnostic, 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 easily be separated 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 comparatively by Mohr (1930), but using only a single species

from each. Much more is needed in the way of comparative morphology

before there can be a logical treatment of the higher categories within

the family.

Scarab larvae are usually C-shaped white grubs, with the head

capsule fairly large, hard, and dark colored. However, many of the

Scarabaeinae and Geotrupinae have a "hump-backed" appearance (Fig. 134).

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 characters are found on the mouthparts

(especially the epipharynx) and the venter of the last abdominal seg-

ment (raster). Near the end of this study an excellent book appeared

summarizing the described North American scarab larvae (Ritcher, 1966).


Biology


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






23
habits and behavior of the organism in relation to both the abiotic and

biotic environment. An attempt is made to summarize our knowledge of

this subject under the individual species treatments, and only brief

generalizations will be made here.

The family as mentioned earlier, can be divided on the basis of

food habits into two major groups which coincide with the morphological

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 primarily 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 investi-

gated (nor has that of any other American coprophage)." The same state-

ment could apply to nearly all members of the family, with the possible

exception of a few economically 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) published an excellent book on the ecology of the sub-

family Aphodiinae, but the work involved only European species.

However, much of the general information on ecology of the dung-feeding

species is applicable to our species. The most recent and most signif-

icant publication on the subject is the volume by Halffter and Matthews

(1966) on "The Natural History of the Dung Beetles of the Subfamily








Scarabaeinae." This work provides an exhaustive review of the literature

of the group, as well as much valuable original data. Howden, in a

series of papers (especially 1955a), has done an outstanding job of

summarizing our knowledge of the North American Geotrupinae. The re-

maining three subfamilies of Laparosticti treated here (Ochodaeinae,

Hybosorinae, Acanthocerinae) are very poorly known biologically. The

food habits of both adults and larvae are unknown.

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 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)). Two species

(Aphodius laevigatus Hald. and Aphodius aegrotus Horn) are found only

with pocket gophers (Geomys spp.). Three species (Onthophagus orpheus

(Panz.), Pseudocanthon perplexus (Lec.), and Ataenius n. sp. #1) have

been found in packrat (Neotoma spp.) nests, although only the latter seems

to be confined to such habitats. One species (Ataenius brevinotus Chapin)

has been found in fox squirrel (Scuirus nigger 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 are myrmecophilous (Euparia castanea Serv. and

Myrmecaphodius proseni Martinez), and occur respectively in the nests of

the fire ants Solenopsis geminata (Fab.) and Solenopsis saevissima Fr.

Smith.








In many of the Scarabaeinae and Geotrupinae the adults provision

the larvae with food in a subterranean chamber (nidification). This

behavior is carried to the ultimate in some species of Copris, in 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 nidification in the

Scarabaeinae, and Howden (1955a) reviewed 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,

but the behavioral significance of sound production has not been investi-

gated. Arrow (1904) described the sound producing organs for a number

of species in the family, and the evolutionary significance of sound pro-

duction of beetles was discussed 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 two 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 dropping, within two hours

after deposition.

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 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

statement 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 is found in the antenna club, and to a much lesser degree

in the maxillary and labial palpi. The two general types of receptors

consist of small conical cells which penetrate the cuticula and end ex-

ternally as pegs (sensilla basiconica), or as short setae (sensilla

trichodea). Another type, in the form of conical pegs arising from small

pits (sensilla coeloconicum), are found on the ultimate segment of the

maxillary palpi.

It is likely that the initial odor stimulation may be a great

distance from the source, but orientation is confined to short distances.

There are probably considerable specific differences in the ability to

perceive and orient toward the food source. In general, stimulation 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 (Fraenkel 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






27
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. It would be especially interesting to learn how those species

associated with certain mammals and ants find their hosts.

Dung beetles are interesting subjects for ecological studies for

several reasons: 1) dung is a rapidly changing micro community 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 numbers, making

the study of populations feasible; 5) a number of species have evolved

unique commensal relationships with other insects and mammals.

In general the dung beetles are seldom attacked by parasitic

Hymenoptera or Diptera. They often harbor mites of many kinds, but this

association is thought to be primarily phoretic. Halffter and Matthews

(1966:171-176) presented a list of the mites associated with various

species of Scarabaeinae. The mites represent several families, chief of

which are the Macrochelidae. These are actively being studied by several

authors, among whom are Costa and Krantz. Most of the material which I

have collected during the course of this study, including several new

species, has been sent to Krantz. In many cases the mites are thought

to be predators on fly larvae (Axtell, 1963) or nematodes (Steward and

Davis, 1967).

There are a number of animals which are predaceous on scarabs, the

most notable being the Surinam toad (Bufo marinus L.) which is successful

in controlling May beetles (Phyllophaga 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 from the stomach of a frog (Rana

sphenocephala Cope). Two skinks (Eumeces egregious (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).

A number of animals such as skunks, opposums, and armadillos undoubtedly

feed on some scarab larvae and adults, but little information is available

on the quantities 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. 138)

contain abundant remains of Dichotomius, Phanaeus, Canthon, Copris, and

Mycotrupes. During this study, Dr. C. T. Collins supplied me with owl

pellets, the data from which will form the basis of a joint paper on the

food habits of this owl in Florida.



Zoogeography


Two things are immediately apparent when one begins to analyze the

distributions of Florida Laparosticti: 1) the high percentage of species

apparently endemic to the state, and 2) the paucity of overlap in both

genera and species with the West Indian fauna.

Of the 106 species and subspecies of Laparosticti recorded from

Florida, there are 22, or slightly more than 20%, not recorded outside

the state. It is likely that some of these will eventually be found in

southern Georgia or Alabama, but they probably have a fairly narrow

range. There are nine additional Florida species that barely range out-

side the state. The endemics are probably the result of many factors,

including insular isolation during past geological periods and unique






29
environmental conditions (e.g. subtropical climate). Only one genus of

Laparosticti (Peltotrupes) is endemic to Florida.

Only 12 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, are in the

subfamily Aphodiinae, and most of them have a fairly wide distribution.

However, one species, Ataenius brevinotus Chapin, is known only from

Cuba and Florida. Another species, Ataenius luteomargo Chapin, known

from 11 islands of the West Indies, is recorded here for the first time

from Key West, Florida, where it is probably a recent introduction.

Psammodius bidens Horn has been recorded from several areas from New

Jersey to Florida and there is a single specimen known from Puerto Rico.

Of the 32 genera of Laparosticti known from Florida, only 11 have

representatives in the West Indies. The subfamilies Geotrupinae,

Hybosorinae, and Ochodaeinae have no genera or species common to both

areas, although the first two subfamilies are represented in both areas

(but by different genera).

Ten of the Florida species, representing seven genera, presumably

have been introduced. One of these (Onthophapus depressus Har.) has also

been found in Georgia and is from Africa. Interestingly, 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 (Blatchley,

1928:13) is valid, it was probably introduced there. Another species

(Myrmecaphodius proseni Martinez), here recorded 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 was introduced into the






30
U. S. probably after 1915, and presumably the beetle was introduced with

its host ant. 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 all 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.

For further details on the distribution and zoogeography, the

reader should consult the discussions under each species in the

systematic account which follows.


Paleontology

There is a single report (Wickham, 1919) of a fossil scarab in

Florida. This is based on an elytral fragment and a smaller fragment

of CoDris inemarginatus 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 undisturbed portions of Stratum

2 (Weigel, 1962), I view this single record with scepticism. 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.






31
Many of the early collections from this site (including the frag-

ments of Copris inemarginatus) were "... carried out without strati-

graphic 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, tortoise, 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 evidence for other insects

at this site, it is unwarranted at this time to attach any great signif-

icance to the two fragments of Copris inemarginatus. It is certainly too

premature to suggest that "... perhaps Copris inemarginatus, a sort of

living fossil, did once utilize the dung of Euus, Mammut, and Mammuthus

which are also found as fossils with it at Vero." (Young, 1959:106).



Economic Importance


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 will be treated later. The Laparosticti, with very

few exceptions, are 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 in-

dividuals constantly 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








incomparably 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 belief. So far as I
can estimate by rough observation, I believe that in certain
active seasons of the year two-thirds of the excrement of this
vast country must be carried by scarabs into the substance of
the soil. Without their valuable aid the land would be an
open sewer. Remember that it supports a teeming population
of some 300 million souls. And, save for the few collected
in the cities, the whole of this great multitude 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 aeration and fertility as a result of this behavior.

Lindquist (1933) measured the amount of dung buried by certain species,

and Bornemissza (1960) suggested that dung beetles could improve

pastures in Australia. There is currently some research being conducted

on the possibilities of introducing dung beetles into Australia and

New Zealand for increasing soil fertility, as well as eliminating

accumulated surface dung.

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






33
unsuitable for complete development of the fly larvae. There have been

several attempts to introduce 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 Hymenolepis 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 hedgehog; Spirocerca lupi (Rudolphi), a

parasite of dog, wolf, jakal, and fox; Physocephalus sexalatus (Molin)

and Gongylonema pulchrum Molin, parasites of a variety of wild and

domestic animals; Macracanthorhynchus 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 para-

sites of man." Stewart and Davis (1967) discussed the "consortisms"

which exist between swine, nematodes, dung beetles, and mites. It

appears that there are many complicated close associations of this sort

which will require detailed studies before any conclusions can be reached.

Although dung beetles serve as intermediate hosts for some verte-

brate parasites, they are also involved in the destruction of eggs,

cysts, and larvae of many parasites. Miller, Chi-Rodriguez, and Nichols

(1961) studied several dung beetles which had ingested human dung
j s








containing eggs of hookworm, Ascaris, and whipworm, and cysts of

Entamoeba coli, Endolimax, and Giardia. No eggs and few cysts were re-

covered from Canthon pilularius (L.) and C. vigilans Lec., and no Ascaris

were found in C. pilularius after the ingestion of embryonated eggs.

Unaltered whipwormnn eggs and cysts, but no hookworm and few Ascaris eggs

were recovered from Phanaeus index MacL. and P. igneus MacL. Our

largest dung beetle, Dichotomius carolinus (L.), regularly contained

and excreted ingested eggs. These authors concluded that "Although dung

beetles may be significant in other ways in dissemination of hookworm,

the species most abundantly attracted to human stools destroy ingested

eggs of this parasite and of Ascaris and do not transport them

internally."













PRELIMINARY CHECKLIST OF FLORIDA SCARABAEIDAE
(Only subfamilies I-VI are treated in this dissertation)


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 Miller
gopheri Hubbard
howdeni Matthews and Halffter
inemarginatus Blatchley
minutes (Drury)

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








Genus 4. Phanaeus MacLeay
igneus igneus MacLeay
igneus floridanus d'l0soufieff
vindex 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 Cartwright
striatulus floridanus Blatchley
subaeneus (Beauvois)
tuberculifrons Harold

Tribe IV. Oniticellini


Genus 1. Oniticellus Serville
*cubiensis Laporte

Subfamily II. .Aphodiinae

Tribe I. Aphodiini

Genus 1. Aphodius Illiger
aegrotus Horn
bicolor Say
campestris Blatchley
crassulus Horn
cuniculus Chevrolat
fimetarius (Linnaeus)
floridanus Robinson
granarius (Linnaeus)
haemorrhoidalis (Linnaeus)
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
proseni Martinez

Genus 3. Ataenius Harold
abditus (Haldeman)
alternatus (Melsheimer)
brevinotus Chapin
cylindrus Horn
erratus Fall
exiguus Brown
fattigi Cartwright
gracilis (Melsheimer)
imbricatus (Melsheimer)
insculptus Horn
linelli Cartwright
luteomargo Chapin
miamii Cartwright
ovatulus Horn
picinus Harold
platensis Blanchard
rudellus Fall
saramari Cartwright
simulator Harold
spretulus (Haldeman)
strigicauda Bates
new species #1
new species #2

Genus 4. Pseudataenius Brown
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. Hybosorinae








Genus 1. Hybosorus MacLeay
illigeri Reiche

Subfamily IV. Ochodaeinae

Genus 1. Ochodaeus Serville
frontalis LeConte
*musculus 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 MacLesy
aeneus MacLeay

Genus 2. Cloeotus Germar
aphodioides (Illiger)
globosus (Say)


Subfamily VII. Melolonthinae








Tribe I. Sericini

Genus 1. Serica MacLeay
aemula Dawson
asper Dawson
atricopilla (Kirby)
delicate 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)
new species #1
new species #2
new species #3

Tribe II. Liparetrini

Genus 1. Hypotrichia LeConte
spissipes LeConte

Tribe III. Melolonthini

Genus 1. Diplotaxis Kirby
bidentata LeConte
frontalis LeConte
languida LeConte
liberty (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 (Schoenherr and Gyllenhal)
latifrons (LeConte)
lota Luginbill
luctuosa (Horn)
marginalis (LeConte)
mariana Fall
micans (Knoch)
okechobea Robinson
ovalis Cartwright
parvidens (LeConte)
profunda (Blanchard)
prununculina (Burmeister)
Ruercus (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.


Genus 1


Pachydemini

.. Gronocarus Schaeffer
autumnalis Schaeffer
multispinosus Howden


Tribe V. Macrodactylini

Genus 1. Macrodactylus Latreille
subspinosus (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
imraculata (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 (Blatchely)

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
Ssquamulosus LeConte






43
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)













SYSTEMATIC ACCOUNT


Key to the Florida Subfamilies of Scarabaeidae

1. Abdominal spiracles situated in the membrane connecting the

dorsal and ventral corneous plates, the last spiracle being

covered by the elytra (Fig. 176); 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------ -- --IAPAROSTICTI-2


it. Abdominal spiracles partly situated in the superior portion

of the ventral plates, the last spiracle usually visible behind

the elytra (Fig. 175); 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---
---------------------------------PLEUROSTICTI-7


2(1). Body globular, capable of contracting into a ball (Fig. 182);

middle and posterior tibiae flattened and dilated (Fig. 163,

165); abdomen with 5 visible ventral segments (Fig. 178);

antennae 10 segmented-------------ACANTHOCERINAE


2'. Body not globular nor capable of contracting into a ball;

middle and posterior tibiae never flattened nor dilated








(Fig. 163, 165); abdomen with 6 visible ventral segments

(Fig. 179); antennae 8-11 segmented- --------- 3

3(2'). Antennae 11-segmented (Fig. 199, 200); mandibles 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. 196, 197,
204) (except for Melanocanthon, in which case the clypeus is

quadridentate); pygidium exposed; mandibles hidden from above
-----SCARABAEINAE


4'. Posterior tibia with two apical spurs (Fig. 180, 181, 201,

202); 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. 13-21); antenna 9-
segmented; epimera of metathorax covered-----APHODIINAE

5'. Mandibles prominent, visible from above (Fig. 139, 140, 152);

antennae 10-segmented; epimera of metathorax covered or

visible -.............. ..-6

6(5'). Body densely pubescent dorsally (Fig. 23); 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. 141-143)--- ---OCHODAEINAE








6'. Body glabrous, shining dorsally (Fig. 22); antenna club with

the first segment hollowed for reception of the second which is

nearly concealed (Fig. 150, 151); color black; both middle

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. 187); not'

more than one pair of abdominal spiracles exposed below edges

of elytra; color yellow-brown to black, never metallic or

green --- ----MEOLONTHINAE


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(80). Front coxae transverse, not prominent; mandibles 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

zreen---.. .....------.... -- ----DYNASTINAE






47
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 metallic and/or green------- -------CETONIINAE













Subfamily SCARABAEINAE (=COPRINAE)
(Fig. 1-12)

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

DIAGNOSIS: Variable in form and size (length 2 to 30 mm), 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. Epimera 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 Scarabaeini).

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, Gynopleurina, 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, although 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 are represented in Florida except the Onitini

(Oriental, Ethiopian and Palearctic), although the single record of the

Oniticellini is doubtful. Only 5 of the 14 subtribes are represented

(i.e. Oniticellina, Dichotomina, Phanaeina, Coprina, and Canthonina) by

12 genera with about 36 species and subspecies.

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 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 necrophagous or saprophagous. Although

practically none of the species are harmful, they have been the subject

of considerable interest for centuries; beginning with their worship by

the ancient Egyptians, through the early classic behavior studies by the

French naturalist Fabre, to the wave of interest of the present day.








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 Matthews (1966). In this paper they have

reviewed the entire literature on the biology of Scarabaeinae, along with

considerable original data on the subject. This publication is by far

the most significant modern contribution to the biology within the family

Scarabaeidae.

This subfamily is the only one of the Scarabaeidae on which there

has been such a review of the biological literature, a situation which

will greatly enhance future work and create considerable interest in

the dung beetles. With such a recent extensive account of the biology,

I will not dwell on the subject further here. However, the reader is

urged to read the above paper for a thorough and enlightening account of

the biology of this fascinating subfamily.


Key to the Florida tribes and genera of Scarabaeinae (-Coprinae)
(Fig. 1-12)

1. Middle and posterior tibiae slender, curved, scarcely enlarged

at the apex (Fig. 197); head and pronotum without horns or

protuberances; elytral striae obsolete, poorly defined- -----
..--.-.......... (Scarabaeini)--4


1'. Middle and posterior tibiae enlarged at the apex (Fig. 196);
head and pronotum often with horns and/or protuberances;

elytral striae obvious or obsolete .-- 2






51
2(1'). Third segment of labial palpi distinct; elytral 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 amm------ 3

3(2'). Antennae 9-segmented; scutellum hidden; several common Florida

species; length 2-8 mm; (Fig. 11)--(Onthophagini)---Onthophagus

3'. Antennae 8-segmented; scutellum small but visible; single

Florida record doubtful; length 5-7 mm; (Fig. 12)----------

.-----..--(Oniticellini)-Oniticellus cubiensis Lap.


4(1). Anterior tarsi absent (Fig. 198); elytral epipleural fold broad;

male with enlarged "hump" on anterior one-third of elytra;

length 20-25 mm; (Fig. 2)------ Deltochilum Z. gibbosum (Fab.)

4'. Anterior tarsi present; elytral epipleural fold very narrow or

absent; neither sex with enlarged hump on elytra; length 2-22 mm

----5

5(4'). Posterior tibia with two terminal spurs; (Fig. 5, 203)-----

--oMelanocanthon

5'. Posterior tibia with a single terminal spur (Fig. 204)------6

6(5'). First segment of posterior tarsus equal to or longer than the

second; surface dull, granular, or shining; color gray to bronze,

if green then dull granulate, not shining; clypeus bidentate or

quadridentate; length 5-22 mmn --- -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 bidentate; length 2-4 mm;

(Fig. 6)----------------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. 7)-- Pseudocanthon perplexus (Leo.)

7'. Base of pygidium margined, clypeus bidentate or quadridentate;

dorsal surface granulate, not shining; color dull gray to green;

length 4-22 mm; (Fig. 3-4)-----8

8(7'). Clypeus bidentate; anterior border of posterior femur margined;
length 11-22 mm; (Fig. 3)---- ---------------Canthon

St. Clypeus quadridentate; anterior border of posterior femur

without margin; length 4-10 mm; (Fig. 4)------Boreocanthon

9(2). Anterior coxae very transverse, not prominent; head and pronotum

without horns or protuberances; color bronze to black, never

bright green; length 5-7 mm; (Fig. 8)--- ---------Ateuchus

9'. Anterior coxae short, conical and prominent; head and pronotum

often with horns or protuberances; color black to bright

metallic green; length 8-29 mm------------------10








10(9'). Anterior tarsi absent in males (Fig. 188), 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. l)-----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

1(10'). Elytral striae seven; clypeus not notched medially; length

20-30 mm; (Fig. 9)--- ----- -Dichotomius carolinus (L.)

11'. Elytral striae eight; clypeus usually notched medially; length

8-15 mm; (Fig. 10) -----:S...Copris


Tribe SCARABAEINI
(Fig. 3-7)

This tribe was divided into seven subtribes by Halffter and

Matthews (1966), but only one (Canthonina) 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 Blackwelder (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 catalogued for the western hemisphere (Vulcano and Pereira,

1964). In this catalogue 39 genera are listed, of which the following

six occur in Florida: Boreocanthon, Canthon, Deltochilum, Glaphyrocan-
thon, and Pseudocanthon.






54
All of the Florida species, except Deltochilum gibbosum Fab., have

been listed in the genus Canthon in previous papers on U. S. species.

Arnett (1962) in his "Beetles of the U. S." does not recognize the

genera Boreocanthon, Glaphyrocanthon, Melanocanthon, 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 origin in the American

tropics, with only a few species entering the U. S. Howden (1966b),

expresses doubt about the validity of certain genera, but he admits 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. The dorsal surface is often granular, and the

elytral striae are poorly defined. The head and pronotum are both with-

out horns or protuberances, although the clypeus is bidentate 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

(Canthon).

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 re-

ported for Canthon pilularius (L.) (Matthews, 1963). These beetles are

familiar to most farm boys and are called "tumble bugs."








Genus DELTOCHILUM Eschscholtz
(Fig. 2)

Deltochilum Eschscholtz 1822:37.

Anamnesis Vigors 1826:510.

Hyboma Serville 1828:352 (not Hyboma Huebner, 1820:200).

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 Eschscholtz 1822:38 (by monotypy).

DIAGNOSIS: Large for the tribe (length to 25 mm, width to 19 am); dull

black; anterior tarsi lacking; middle and posterior tibiae long, slender,

the posterior ones bent inward abruptly at the middle, scarcely enlarged

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

longitudinal) carinae at this point. Humeral carina short and longi-

tudinal. 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), Euhyboma (1),

Telhyboma (1), Hybomidium (7), Parahyboma (2), Rubrohyboma (1),

Deltohyboma (43), and Deltochilum (3). Howden (1966b) added two new

subspecies, one new species, and synonymized one species to bring the

'total to 75 species and subspecies currently recognized.

Our single species, _D. gibbosum (Fab.), is the type of the subgenus

Hybomidium Shipp (1897:195). This name was proposed to replace Hyboma

Serville (1828:352) which was preoccupied by Hyboma Huebner (1820:200)

in the Lepidoptera. Paulian (1938:259) subsequently described the new

subgenus Tetraodontides with D. gibbosum as the type species, although

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 Hybomidium, our species

becomes Deltochilum (Hybomidium)gibbosum (Fab.)

DISTRIBUTION & ZOOGEOGRAPHY: The genus is almost exclusively Neotropical,

with only two species entering the United States. Of these two, D.

scabriusculum Bates is known from Brownsville, 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 primarily 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 and

animal material. 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 decaying cantaloupe in the three forested habitats

investigated, but none were 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 de-

composition. 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. gibbosum was attracted to dead fish

within one day after the frest bait was deposited, but cantaloupe was

not attractive until after the 7th day when the flesh was "... very soft,






58

much liquid." My own experiences in Mexico and Central America indicate

that, although the habitats are variable, specimens are rarely encountered

in open pastures. Gibson (In Howden, 1966b:738) found D. scabriusculum

montanum Howden active around horse dung, but none were 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.

197) 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 un-

like that of the smooth, round ball of Canthon. They are most frequently

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 excluding 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 Coprinae. The most use-

ful taxonomic character for separating them from other genera appears to

be the setal pattern of the venter of the last abdominal segment. Larval
development 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,

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 (Fabricius)
(Fig. 2)

Scarabaeus gibbosus Fabricius 1775:28.

Copris gibbosus (Fab.), Olivier 1790:141, 172.
Ateuchus gibbosus (Fab.), Fabricius 1801:57.
Hyboma gibbosa (Fab.), Serville 1828:353.
Deltochilum gibbosum (Fab), Burmeister 1848:134.
Deltachilum gibbosum (Fab.), LeConte 1863b:36 (misspelling).

Deltochilum g. gibbosum (Fab.), Bates 1887:36.

DIAGNOSIS: Easily distinguished from other Florida Canthonini by the

lack of anterior tarsi, the broad epipleural 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 south-

eastern U. S. Howden (1966b:736) stated, "... the species is represented

by a complex of related forms extending through Mexico and Central America

into South America. Several seemingly disjunct populations show constant

differences, but, because 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 relationship, and partly to indicate the possibility

of interbreeding." He recognized two additional subspecies: sublaeve

Bates from Mexico and panamensis Howden from Panama.


DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 33). Recorded from Alabama,

Georgia, Louisiana, Mississippi, North Carolina, South Carolina,

Tennessee, Texas, and I have also seen specimens from southeastern

Kentucky (new state record). In Florida it occurs from Pensacola to Big

Pine Key, the spotty records probably reflecting its secretive habits,

and it probably occurs throughout the state.


BIOLOGY: (see remarks under the genus). This species 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: 55, of which 40 were from 16 Florida localities

(for complete data see appendix 1).

SELECTED REFERENCES: Angell, 1913:169; Blatchley, 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 Martinez, 1956:125; Vulcano

and Pereira, 1964:648-649; Walker, 1957:Table 5,7, Fig. 6, 9-11.


Genus CANTHON Hoffmannsegg
(Fg. 3)

Canthon Hoffmannsegg 1817:38.

Coprobius Latreille 1829:535.

Coeloscelis Reiche 1841:213.

TYPE SPECIES: Scarabaeus pilularius Linnaeus 1758, by subsequent

designation of Paulian (1938-39:22).

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 mono-

graphed for North America (Halffter, 1961) and catalogued for the world

(Vulcano 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 Glaphyro-

canthon, Pseudocanthon, Boreocanthon, and Melanocanthon. Numerous

other genera have also been split from the old genus Canthon, especially

for the South American 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).

Our Florida species are similar in appearance and often difficult

to determine without comparative material. 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 distin-

guishing many of the South American forms. No subspecies are recognized

for the U. S. species, although several have been described for Mexican

and Central American species (Halffter, 1961). Our species vary pri-

marily 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).

DISTRIBUTION & ZOOGEOGRAPHY: Nearly all 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)

lists a single specimen from Key West. Of the remaining 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 fann boy. All of our species are primarily coprophagous

and feed commonly 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 grassland biome. Only one of our species (pilularius)








has been studied in any detail (Matthews, 1963). Only general behavior,

probably common to all of our species, 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 behind 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. Comstock (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 nocturnal, possessing enlarged eyes,

and it is the only U. S. species attracted to light.







SELECTED REFERENCES: Balthasar, 1939; Blanchard, 1885; Brown, 1928a;

Halffter, 1961; Harold, 1868c; Paulian, 1939; Pereira and Martinez,

1956; Robinson, 1948b; Schmidt, 1922b; Vulcano and Pereira, 1964.


Key to the Florida species of Canthon

1. Eyes large for the genus (Fig. 189); larger (length 15-22 mm);

color uniform black; often attracted to light-----vigilans Lec.

i'. Eyes normal for the genus (Fig. 190); smaller (length 11-18 mm);

color usually bronze, bluish or greenish, rarely black; never

attracted to light------------------2

2(11). Head, pronotum, and pygidium with uniform minute granules,

never with enlarged granules as are present on the elytra;

uniformly 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; color variable from bronze to bluish,

greenish, or dull matte gray; common Florida species--------

.......--- ilularius (L.)


Canthon chalcites (Haldeman)

Coprobius chalcites Dejean 1836:151 (nomen nudum).

Coprobius chalcites Haldeman 1843:304.

Canthon chalcites Hald., LeConte 1859a:10.

Canthon chalcides Hald., LeConte 1863b:36 (misspelling).






66
DIAGNOSIS: Easily distinguished from the two other Florida species by

the lack of coarse granules on the head, pronotum, and pygidium like

those of the elytra. It differs further from vigilans Lec. in the

narrower eyes as in pilularius (L.). It differs further from pilularius

by generally larger size (length 13-21 mm) and uniformly bronze (or

rarely black) color. Anterior tibial spur sexually dimorphic (Fig.

193-195).

TAXONOMIC NOTES: Harold (1868c) synonymized this species under

pilularius (as laevis), but he was clearly in error in doing so, as was

pointed out by Horn (1870a). Although closely related to that species,

the differences in granulation are very apparent on comparison of the

two. Occasionally 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 shining spots

without any height. He further postulates that these two forms "...

may prove to be subspecific 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

Nebraska, it is rarely collected elsewhere. Until this material is

available in numbers, a proper evaluation of the variability cannot be

made.








DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 34). The type is from

"Missouri." Halffter (1961:300) recorded it from Alabama, Florida,

Georgia, Illinois, Kansas, Kentucky, Louisiana, Massachussetts, 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 Miami. It was not recorded in the "Scarabaeidae of Florida"

by Blatchley (1927-30).

BIOLOGY: Practically nothing has been published on the habits of this

species. Presumably it is a dung feeder and rolls balls similar to 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 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 five 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) 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).








SELECTED REFERENCES: Blanchard, 1885:166; Dillon and Dillon, 1961:
508, Pl. 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. 3)

Scarabaeus pilularius Catesby 1731-1743: Pl. 11 (Pre-Linnean name).

Scarabaeus pilularius Linnaeus 1758:349.

Scarabaeus laevis Drury 1770:79, Pl. 35, Fig. 7.

Scarabaeus hudsonias Forster 1771:24.

Scarabseus 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. 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. 190).

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 lists the

locality as "America." However, he also cites references to pilularius

by Pliny and Aristotle (presumably European and thus a member of

Gymnopleurus and not Canthon). Numerous 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, Florida, and later synonymized by Halffter (1961).

I am personally convinced that the subspecies was based on mislabeled

specimens, and imitator, in any of its variations, does not occur in

Florida. The types of floridanus 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 labeled specimen. Since these specimens do not carry a

specific date nor a field number, their status is immediately 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 mislabelled 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 material."






70
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 forms.


DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 35). It apparently occupies

nearly the entire area east of the Rocky Mountains. Previous records

from Mexico are probably all referrable to the closely related imitator.

The record of Blanchard (1885:166) for "S. Cal." is probably also in-

correct since no subsequent specimens 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

"America," but Robinson (1948b:93) listed the type locality as "New

York."


BIOLOGY: As the common name "tumble bug" implies, 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 con-

structing the balls. Since this presumably occurred in Oklahoma, this








record could refer to either pilularius or imitator. The life history

was described by Lindquist (1935), Cooper (1938a), Ritcher (1945), and

Miller (1954). The behavior 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. Matthews 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 provisioning the larva. For

a general discussion of ball rolling in dung beetles see the discussion

on nidification 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 have been described for

comparison. It can be separated from the other known larvae of the sub-

family 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 abdominal segment with two inconspicuous patches of very short setae.

Although there is probably more published information on this species

than any other dung beetle, there is much to be learned about behavior

and morphological variation.

MORPHOLOGY: The internal anatomy was studied by Cooper (1938b),

and it is this study which is the basis for most generalizations on dung






72
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

exceptionally 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 respiratory system

shows no air sacs arising from abdominal tracheae, but seven pairs arise

from the metathoracic spiracles and form a group candad to the heavily

muscled thoracic region. Air sacs from the mesothoracic spiracles extend

forward into the head.

The external morphology was described by Mohr (1.930), and Halffter

(1961). The mouthparts were described in detail by Hardenburg (1907)

and Miller (1961).

SPECIMENS EXAMINED: Over 500, of which 246 were from Florida (for

complete data see appendix 2).

SELECTED REFERENCES: The more important references 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

Canthon vigilans LeConte 1858:16.

DIAGNOSIS: Distinguished from the two other Florida species by the

greater width of the eye as seen dorsally (Fig. 189). 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 exclusively nocturnal. It reaches the maxinmm

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. Additional

specimens will be required to determine the full extent of this variation.

DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 36). Originally described from

Georgia, Missouri, and Texas, without specific designation of a type

locality. Robinson (1948b: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, Illinois, Kansas, Massachussetts, Michigan, Missouri,

Mississippi, New Jersey, New York, North Carolina, Texas, and Virginia.

I have also seen specimens from Kentucky 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,

in nearly all cases 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 deaphic tolerances. Miller (1954:Table 1-2) found this species

attracted to traps containing human feces at night in Georgia. The

immature stages are unknown.

SPECIMENS EXAMINED: About 100, of which 63 were from 15 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.


Genus BOREOCANTHON Halffter
(Fig. 4)

Boreocanthon Halffter 1958:208-209.

TYPE SPECIES: Canthon ebenus (Say), by original designation.

DIAGNOSIS: Similar in general appearance to Canthon. 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. It is easily separated from

Melanocanthon, which it closely resembles superficially, by the single

spur on the posterior tibia.

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 characters

given in the key.


DISTRIBUTION & ZOOGEOGRAPHY: The genus, 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 coprophagous. 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 enlarged granules

scattered over the minutely pebbled surface; anterior tibial spur

of the male elongate, pointed, not bifurcate; larger (length

7-10 mm)------- -- -depressipennis Lec.


lt. 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.


Boreocanthon depressipennis (LeConte)
(Fig. 4)

Coprobius depressipennis Dejean 1836:152 (nomen nudum).

Canthon depressipennis LeConte 1859a:11.

Canthon depressipenne Lec., Blackwelder 1944:199.

Boreocanthon depressipennis (Lee.), Halffter 1958:208.

DIAGNOSIS: Easily distinguished from our only other species by the

characters in the key. In addition specimens sometimes have green or

blue reflections. The male genitalia are 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 con-

fusion or synonyms.

DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 37). Originally described from

Kansas, although Robinson (1948b:92) 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) reported it from Suwanee Springs, and

Castle and Laurent (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 pan-

handle 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 September.

Nothing else seems to have been published on its habits, and the

immature stages are unknown.

SPECIMENS EXAMINED: 221 from 23 Florida localities (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

punctate (often only visible at an oblique view at high [90X] magnifi-

cation). 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

LeContets unique, it must be remembered that both species have been de-

scribed from single specimens."

There is some variation in size (length 4-6 mm) and in the coarse-

ness 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 differences.

DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 38). It was originally described

from "America septentrionali," and Kansas was the type locality for the

synonym 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);

Alabama (Loding, 1945:98); Oklahoma, Texas, Utah, and Virginia (Robinson,

1948b:90). I have also seen specimens from Arizona (new state record).






79
It was first recorded from Florida (Enterprise) by Schwarz (1878:

449). Blatchley (1918:54) added Crescent City and St. Augustine, and

later (1928:61) included Centerville, St. Mary, Marion Co., and Dunedin.

My records include nearly the entire peninsula, but there are no records

west of Jefferson Co. However, this is probably an artifact of collecting,

since it was recorded from Mobile Co., Alabama (Loding, 1945).

BIOLOGY: My experience has been that this species is rather uncommon

in Florida. However, this observation is based on collecting in cow

dung in pastures. It appears to be more abundant in sandy, wooded sit-

uations. 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: 60, of which 47 were from 21 Florida localities

(for complete data see appendix 5).

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.

Genus MELANOCANTHON
(Fig. 5)


Melanocanthon Halffter 1958:210-211.







TYPE SPECIES: Canthon bispinatus Robinson (by original designation).


DIAGNOSIS: Superficially similar to Canthon and Boreocanthon, but

differing from both, and from all other members of the subfamily Scara-

baeinae, in possessing two spurs on the posterior tibiae. Clypeus

quadridentate, 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).


DISTRIBUTION & ZOOGEOGRAPHY: The genus is 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 Plains species.

BIOLOGY: The species are usually rarely collected. For instance, when

Robinson (1941) revised the group he saw only 86 specimens in the five

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, 1958:210-2100, Fig. 7-11; Robinson,

1941:127-130, Fig. 1-8; Vulcano and Pereira, 1964:594-595.



Key to Florida species of Melanocanthon


1. Pronotal granules reduced to shining spots, none in relief;

head and pronotum noticeably punctate-------- ----------
----------punctaticollis (Schaeffer)


1'. Pronotal granules in relief, 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 noticeable--------ranulifer (Schmidt)


2'. Granules of dorsal surface less dense, less raised; punctures

of the head as noticeable and as evenly distributed as the

granules ---------------------- bispinatus (Robinson)


Melanocanthon bispinatus (Robinson)

Canthon bispinatus Robinson 1941:128-129, Fig. 2-3.

Melanocanthon bispinatus (Robinson), Halffter 1958:210-211.


DIAGNOSIS: Although the male genitalia are very; distinctive, it is

otherwise often difficult to separate from granulifer, without compar-

ative material. In general the granules are less pronounced and often

reduced to elongate, shining, black spots in the antero-median are of

the pronotum. The punctures of head and pronotum, although perhaps no

more numerous, are more noticeable because of the greater space between

the granules.






82
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. It

was made the type of the genus Melanocanthon by Halffter (1958:210).


DISTRIBUTION & ZOOGEOGRAPHY: (map, Fig. 39). The type locality is

Warren Grove, Burlington Co., New Jersey. Paratypes were recorded from

the following states: Alabama, Florida, Georgia, New Jersey, North

Carolina, Rhode Island, South Carolina, and Virginia.

In Florida it has been reported from DeFuniak Springs and

Enterprise (Robinson, 1941:129). My records add seven 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. nigricornis 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 specimen rolling a ball

and this particular ball was composed of deer excrement." I can add only

that several specimens were taken in malt bait traps in Florida.

Specimens have been taken from March through September, but in Florida

they seem to be more abundant in April. The immature stages are unknown.




University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs