ARTHROPODS OF FLORIDA
and Neighboring Land Areas

VOLUME 13

THE SCARAB BEETLES OF FLORIDA

(Coleoptera: Scarabaeidae) Part II.
The May or June Beetles
(genus Phyllophaga)

by
Robert E. Woodruff
and
Brenda M. Beck

v.3

Florida DD'-me- of AWricum and C-sumr Srvic=s
Doyl C=, C'-ommio
Di43 n.ofPLa26Indus
Rlchad Oaska~Lb, DlWror
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v.13

ARTHROPODS OF FLORIDA
and Neighboring Land Areas
VOLUME 13

THE SCARAB BEETLES OF FLORIDA

(Coleoptera: Scarabaeidae) Part II.
The May or June Beetles
(genus Phyllophaga)

by
Robert E. Woodruff
and
Brenda M. Beck

Typical position of May beetles, feeding and mating
at night; female on the left [Phyllophaga quercus
(Knoch)]. Photo by Jeff Lotz.

Florida Department of Agriculture and Consumer Services
Doyle Conner, Commissioner
Division of Plant Industry
Richard Gaskalla, Director
P.O. Box 1269
Gainesville, Florida 32602

Contibin No. 716, Buu of Enmomology
Release Dae: Oc. 1,989
jL'RlITY OF FLORIDA UIRAIS

SCIEWC

This public document was promulgated at a cost of $38,214.00 or $26.00 per copy. It makes available to
all interested persons the results of arthropod faunal studies, emphasizing Florida and the Circum-
Caribbean Region. PI89T-37

ISSN: 0066-8036

DEDICATION
It is with great pleasure that we dedicate this
volume to Dr. Milton W. Sanderson, the dean of
North American Phyllophaga specialists for over
50 years. He was the moving force which
enabled this study to be completed, and he has
always been willing to share his phenomenal
knowledge with younger workers. His monu-
mental collection, accumulated during his tenure
at the Illinois Natural History Survey, will be the
mecca to which all future specialists must go.
His personal friendship to the senior author
resulted in the donation of his library, manu-
scripts, many new species, and other pertinent
materials. Without his stimulation and continued
support, this volume would not have been
completed.

TABLE OF CONTENTS

Title Page ..................................................................................................................... i
Dedication .................................................................... ............................................... ii
Table of Contents .............................................................................................................. iii
Foreword ...................................................................................................................... iv-vi
List of Tables .................................................................................................................... vi
Preface .........................................................................................................................
Acknowledgm ents .......................................................................................................... 1-3
Abstract ......................................................................................................................... 3
Plates 1-35 (fig. 1-410) ................................................ ........................................... 4-38
Alphabetical Checklist, with Figures and Maps. Table 1. ........................................ 39-40
Introduction ...................................................................................................................... 41
Form at of Presentation .................................................................................................... 41
Historical Resum 6 ................................................................................................ ..... 44
Chronological List of Florida Phyllophaga descriptions. Table 2. ........................... 44
M methods and M materials .................................................................................................... 45
Collecting Techniques ................................................................................................... 46
General Account of the genus Phyllophaga ...................................................... ............. 46
Taxonom y ................................... ................. .......... ....................... ..................... 47
Arrangement of Florida Phyllophaga by groups. Table 3 .............................................. 48
M orphology .................................................................................................................... 49
Biology & Behavior................................................................................................. 55
Plant H costs ............................................................................................................. 56
Im m ature Stages ................................................................................................... 61-64
Known Life Cycle Summary of Florida Phyllophaga. Table 4 ............................. 56
Alphabetical List of references to Known Larvae. Table 5 ................................. .. 65
Key to Known Larvae of Florida Phyllophaga .................................................... 63-64
D distribution and Zoogeography ................................................................................ 65-66
Economy ic Im portance ......................................................................................... 66-67
N natural Enem ies ...................................................................................................... 67-70
Parasites ....... ......................................................................... ............................. 68-69
Predators ...................................................................................................................... 69
M icroorganism s .................................................................................................... 69-70
Questionable and Potential Florida Records .............................................................. 70-73
Misspelled Use of Florida Phyllophaga Names. Table 6 ............................................ 72
Checklist of Florida Species (including synonyms). Table 7. ..................................... 74-75
Selected Taxonomic Characters of Florida Phyllophaga. Table 8. ........................... 76-77
Key to Adults of Florida Phyllophaga .................................................................... 78-83
System atic Accounts by Species .................................... ......... ........ ..................84-170
Bibliography ................................................................................... ...................... 171-183
Appendices 1-39 (Specimen Data), p.184-217; 49 (List of Figures), p.218-223 ........ 184-223
Index ........................................................................................................................ 224-225

FOREWORD

This is the Part II of The Scarab Beetles of
Florida, initiated in 1973 by Dr. Robert E. Woodruff
with Volume 8 of Arthropods of Florida and
Neighboring Land Areas, (Coleoptera: Scarabaeidae)
Part I. The Laparostici (Subfamilies: Scarabaeinae,
Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae,
Acanthocerinae). The high standards of accuracy and
attention to detail are continued in this volume. The
difficult task of preparing a treatment of the subject that
is understandable to the novice and at the same time a
scholarly and comprehensive presentation for other
systematics authorities is accomplished.

This volume is based on more than 30 years of
extensive collecting in many parts of Florida in all
seasons of the year by the senior author, using a variety
of collecting techniques. As in the first volume, virtu-
ally all known collection records for the group studied
are given, and questionable and potential Florida rec-
ords are considered. Over 100,000 specimens were
examined.

In their treatment of the large and difficult to
understand genus Phyllophaga, the authors have made
extensive use of genitalia as the most reliable means of
species identification. Genitalia of all known Florida
species and those of some closely related species are
exceptionally well illustrated through the use of scan-
ning electron microscope photographs of the highest
quality. An extensive bibliography documents infor-
mation presented in this volume and presents refer-
ences for anyone who wants to study the subject further,
although the present volume gives a comprehensive
treatment of whatis known about Florida Phyllophaga.
The present faunal study, like that treated in Part I,
is provided primarily as a manual to assist in the iden-
tification of the Florida species. Specific identification
of an organism is essential to understanding the role it
plays in the environment and in determining the course
of action for controlling those species that become a
problem to humans. The name is the "key" to the
published literature and is essential before controls are
attempted. Illustrated keys for the identification of the
Florida species are provided along with an adequate
description of each species.

Dr. Robert Eugene Woodruff or "Bob" as he is
called by his friends and associates, was born in Ken-
nard, Ohio, on 20 July 1933, son of Marvin C. and
Thelma B. Woodruff. On 27 June 1954 he and Nina
Evelyn Gunsaulies were married in Urbana, Ohio,
where Bob's parents still reside. They have 2 children,
Kris Eugene Woodruff, 29, now living in San Juan,
Puerto Rico, and Cheri Ellen Taylor, 33, now residing
in Silver Spring Shores, Florida. Bob and Evelyn have
2 grandchildren. Bob was educated in the public
schools of Ohio. From early childhood he expressed an

interest in the natural sciences. In 1950 and 1951 he
entered the Junior Science Fairs of the Ohio Academy
of Science, receiving superior awards both years and a
scholarship to any of the 10 state universities. Follow-
ing graduation from Urbana High School in 1951, he
enrolled at Wabash College in Crawfordsville, Indiana.
During 1951-1952 he served as Curator of the insect
collection at Wabash College. The following year he
transferred to Ohio State University where he received
the Bachelor of Science degree with a major in ento-
mology in 1956.

From 1952 to 1955 he was an assistant in the De-
partment of Natural History of the Ohio State Archaeo-
logical and Historical Society Museum where he gained
valuable experience and training under Dr. Edward S.
Thomas and Mr. Robert Goslin. From 1955 to 1957 he
held a graduate assistantship in the Department of
Zoology and Entomology at Ohio State University,
under Professor Josef N. Knull, Curator of the Entomo-
logical Museum. This experience led him to specialize
on the beetle family, Scarabaeidae. Prof. Knull intro-
duced him to the "scratch board" technique of beetle
drawings which he has used in most of his publications,
including 32 habitus drawings in Part I of The Scarab
Beetles of Florida, published in 1973.

From 1957 to 1958 he was employed as a medical
entomologist with the Kentucky State Health Depart-
ment, in Louisville, working on St. Louis Encephalitis
and related mosquito projects. He attended special
courses of instruction on insects of medical importance
at the U. S. Public Health Service, Communicable
Disease Center, in Atlanta, and at the Tennessee Valley
Authority, Wilson Dam, Alabama. In March 1958 he
joined the staff of entomologists at the State Plant
Board of Florida (now Division of Plant Industry,
Florida DepartmentofAgriculture and Consumer Serv-
ices). He was initially employed as the "Survey
Entomologist" to coordinate the federal-state Coop-
erative Economic Insect Survey, in Florida, which he
fulfilled until 1963. The responsibilities included field
surveys, identifications, and reports for pests of turf,
field crops, vegetables, fruit, pastures, and ornamental
plants.

From 1963 more emphasis was on detection of
foreign pests and primary identification for beetles
(Coleoptera) and grasshoppers (Orthoptera). Biologi-
cal control and pest management have been investi-
gated with pests introduced from Latin America. Re-
sponsibilities included curating the Coleoptera and
Orthoptera for the Florida State Collection of Arthro-
pods, now the fifth largest arthropod collection in the
United States, and the development of the entomologi-
cal portion of the DPI library. Consultancies were
interspersed with this position from 1978. Retirement

after 30 years of service for the State of Florida was
effective 31 July 1988. He continues to serve the
program as an Emeritus Entomologist.

In September 1963 Bob entered the Graduate School
of the University of Florida, from which he received the
Doctor of Philosophy degree with a major in entomol-
ogy in 1967. As a part of his studies he attended a
session of the Organization for Tropical Studies at the
University of Costa Rica. His research has taken him to
much of the U. S. and the following countries where he
has collected and studied specimens: Antigua, Argen-
tina, Australia, Barbados, Bolivia, Brazil, Colombia,
Costa Rica, Cuba, Dominican Republic, El Salvador,
Fiji, Grenada, Guadeloupe, Guatemala, Haiti, Hondu-
ras, Jamaica, Mexico, Montserrat, Nevis Island, Nica-
ragua, Paraguary, Peru, Puerto Rico, St. Kitts, St.
Lucia, St. Vincent, Trinidad, Uruguay, and Venezuela.
He has received grants and contracts from the Ohio
Academy of Sciences, United States Public Health
Service, National Science Foundation, Smithsonian In-
stitution, U. S. Department of Agriculture, Australian
Academy of Sciences, Florida State Museum (since
July 1988 the Florida Museum of Natural History),
Illinois Natural History Survey. In 1988 he held a
consultancy with FAO (United Nations) in Barbados,
St. Lucia, and Grenada; and Agro Delta in the Domini-
can Republic. In 1989 he held a consultancy with Texas
A. & M. University in College Station, Texas and
Dallas, Texas. He is currently consulting with United
Nations, FAO and has a 6-month Fulbright Fellowship
for research in Paraguay.

Societies in which Dr. Woodruff holds member-
ship include: Association for Tropical Biology, Cole-
opterists Society (President, 1978; Editor, 1972-1977),
Entomological Society of America, Florida Entomo-
logical Society (Associate Editor, 1971-1977), Gamma
Sigma Delta, Phi Kappa Psi, Sigma Xi, Sociedad Mexi-
cana de Entomologia, Society of Systematic Zoolo-
gists, and Gainesville (Florida) Gem & Mineral Soci-
ety. He is an Adjunct Professor, Department of Latin
American Studies, University of Florida, Adjunct Pro-
fessor, Department of Entomology & Nematology,
University of Florida; Courtesy Associate Professor,
Department of Entomology & Structural Pest Control,
Florida A. & M. University, in Tallahassee; Adjunct
Curator, Department of Natural Science, Florida Mu-
seum of Natural History (since 1972); Research Asso-
ciate, Museo Nacional de Historia Natural, Santo Dom-
ingo, Dominican Republic; and Collaborator, USDA,
APHIS (1960 to present). In 1986 he received an award
(with H. A. Denmark, F. W. Mead, and H. V.
Weems, Jr.) from the Insect Behavioral Ecology unit
for taxonomic service, Florida Entomological Soci-
ety. Since 1983 he has served on the Board of Direc-
tors, (Chairman) Caribbean Center for Scientific
Research (non-profit). Since 1986 he has served on the
Board of Directors of PhytoTechnica Floridana, con-
sulting corporation. He is a member of the Board of
Directors of the "North American Beetle Fauna Proj-
ect". In 1986-1987 he served as the first President

of the Center for Systematic Entomology, Inc. He
received an Honorary Award from the Eastern
Federation of Mineral &Lapidary Societies, to select
scholarships by the American Federation of Mineral
Societies Scholarship Foundation for 1989.

Dr. Woodruff has served as Editor of The Journal
of the Newell Entomological Society (President, 1965-
1966); Editor, Coleopterist's Newsletter, 1970; Asso-
ciate Editor, Florida Entomologist, 1969-1975; Editor,
Coleopterist's Bulletin, 1971-1975; Associate Editor,
1975-1982; Managing Editor, Insect World Digest,
1971-1974; on Council of Biological Editors, 1972-78;
member, Editorial Board, Colemania, Indian Journal
of Taxonomic Entomology, 1981-present; member
Editorial Board, Insect Mundi, 1985-present. He has
attended more than 30 national and international scien-
tific meetings, including the 14th International Con-
gress of Entomology, in Washington, D. C. (for which
he designed the logo). In 1964 he attended two months
of training attheOrganization for Tropical Studies, San
Jose, Costa Rica. In 1971 he attended the Summer
Institute for Systematics, Smithsonian Institution. He
has presented papers at meetings of the Entomological
Society of America, National PestControl Association,
Ohio Academy of Sciences, Sociedad Mexicana de
Entomologia, International Congress of Entomology,
Florida Entomological Society, and other scientific
organizations. He has published over 150 papers, pri-
marily in the field of taxonomic and economic ento-
mology.

Dr. Woodruff is listed in American Men of Science,
Directory of Zoological Taxonomists, The Naturalists
Directory, International Scholars Directory,
Personalities of the South, Who's Who in the South
and Southeast, and Directory of North American
Entomologists. He is a Methodist. His hobbies include
archaeology, paleontology, scientific illustration, tropi-
cal biology, lapidary, jewelry, and music (trumpet,
guitar, voice). He has served as merit badge counsellor
for the Boy Scouts in all natural history subjects.

Brenda Miller Beck was born in Key West, Florida
on 23 October 1948, daughter of George P. and Bonnie
S. Miller. Brenda and Dennis A. Beck were married 1
July 1970 in Gainesville, Florida. They have one
daughter, Jessica Denise, age 5. Brenda received her
elementary and high school education in the public
schools of Florida, Oklahoma, and Texas, graduating
from Gainesville High School in 1967. Brenda worked
at the University of Florida during the summer prior to
her last year of high school. In March 1974 she received
the Associate of Arts degree from Santa Fe Commu-
nity College, in Gainesville, Florida. She began work
with the Division of Plant Industry in 1968 and has
worked continuously in the Bureau of Entomology for
21 years. Currently a Laboratory Technologist IV,
Brenda has worked extensively with Coleoptera, par-
ticularly in the preparation of genitalia ofscarabaeid
beetles for study, involving scanning electron photo-
microscopy, and identification of Scarabaeidae and

tera. For many years, under the tutelage of Dr. Woodruff,
and also for the past year under Dr. Michael Thomas,
Brenda has served as the principal, active curator of
the Coleoptera and Orthoptera collections of the
Florida State Collection of Arthropods. Shealsohas
worked extensively in the curating of the large
alcohol-preserved, compactor-stored collections of
the FSCA, and the computerization of collection data.
In recent years she has worked also with Dr. Howard
Weems on the annual summary of all donations to the
FSCA, the computing of estimated values of these
donations, and preparation of several thousand
formal letters of acknowledgment which have been
sent to donors. Working with Dr. Woodruff, Brenda
has acquired a considerable knowledge of the identifi-
cation of Scarabaeidae and other Coleoptera. Her
current research interest is centered on the Phylloph-
aga of the United States and the West Indies. In

February 1989, at the request of Dr. Robert L.
Crocker of Texas A. & M. University, Brenda spent a
week at Texas A. & M. training personnel on the
extraction of Scarabaeidae genitalia and their prepara-
tion for study. Brenda was raised as a Baptist Her
hobbies are raising horses and dogs and working with
plants. She also enjoys writing poetry and some fiction.

Bureau of Entomology
Division of Plant Industry
Florida Department of Agriculture
and Consumer Services

1 August 1989

Howard V. Weems, Jr.
Editor

List of Tables

1. Alphabetical checklist with figures and maps ...................................................................................... 39-40
2. Chronological list of Florida Phyllophaga descriptions .................................................................. 44
3. Arrangement of Florida Phyllophaga according to groups by various authors ........................................ 48
4. Known life cycle summary of Florida Phyllophaga. ........................................................................ 56
5. Alphabetical list and references to known larvae of Florida Phyllophaga............................................. 65
6. List of misspelled use of Florida Phyllophaga names ................................................................... 72
7. Annotated alphabetical checklist of names used for Florida Phyllophaga........................................... 74-75
8. Selected taxonomic characters of Florida Phyllophaga ..................................................................... 76-77

PREFACE

When Part I of this series on the "Scarab Beetles
of Florida" was published (Woodruff, 1973), it was
anticipated that succeeding parts would be completed
soon thereafter. The press of routine duties, consider-
able travel and field work, and the volume of speci-
mens have each contributed to this 16-year delay.
The senior author's retirement in August, 1988,
provided the impetus and opportunity to complete the
manuscript. The delay allowed the inclusion of many
more specimens and permitted the illustration of the
genitalia and other taxonomic characters by use of the
scanning electron microscope. We believe it was
worth the wait, and the resulting publication should be
more useful for identifying the Florida fauna of these
interesting insects.
This part is co-authored, the work and responsi-
bilities being equally shared, except for the final
writing. The authors have jointly worked for over 50
years with the Florida Department of Agriculture,
during which varying amounts of time were spent
preparing specimens and data for this eventual publi-
cation. We have also prepared many of the specimens
and recorded much of the data for the final part (III),
which will include the remainder of the Melolonthinae
and the subfamilies Rutelinae, Dynastinae, and Ce-
toniinae. We anticipate completion of Part III without
the attendent delays previously encountered.
As with Part I, emphasis was placed on preparing
a faunal study to enable the user to identify specimens
and then to learn what is known about that species. It
is therefore a compilation of original data, integrated
with the existing literature on the Florida species.
Such faunal surveys are useful to a wide audience,
including homeowners, biologists, biogeographers,
ecologists, entomologists, pest control operators, and
environmentalists.
Permanent preservation of vouchered specimens
is an important part of any such study. The Florida
State Collection of Arthropods* is the primary reposi-

*The Florida State Collection of Arthropods is com-
posed of several collections which were previously main-
tained as separate: Univ. Florida, Agr. Exp. Sta.; Univ.
Florida, Dept. Entomology and Nematology; 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 abused by the Division
of Plant Industry, Florida Dept. Agr., Gainesville, Florida
32602.

tory for our specimens, and it is now the second best
collection of this genus in the World (behind the
Illinois Natural History Survey; see Woodruff, 1987).
Many times during this study we were stymied and
unable to solve problems, because previously reported
specimens were not documented and properly vouch-
ered in a permanent public collection.
Our emphasis on providing an identification manual
required that some traditional elements were short-
ened by time and space restrictions. Although the
species of Phyllophaga are externally similar, their
distinctive genitalia provide exceptional characters for
identification. As a result of this, long verbal descrip-
tions are both superfluous and misleading. We thus
chose to expend our time and energies in preparing the
genitalia illustrations from scanning electron micro-
scope photographs (Plates 1-32, fig. 1-378).
We believe that these illustrations should permit
easy comparison and positive identification, far better
than words could convey. Ironically, just 100 years
ago, Smith (1889b:485) first used these structures with
the following comments: "No words could accurately
describe their peculiar turning and twistings. I shall
not undertake verbal descriptions of these parts, but
prefer to let my figures answer most questions."
We hope that this volume will stimulate collectors
and students to pursue the many gaps in ourknowledge
of a dominant element of our diverse Florida insect
fauna. If it accomplishes these goals, we will be repaid
adequately for the efforts expended in its preparation.

ACKNOWLEDGMENTS

As employees of the Division of Plant Industry,
Florida Department of Agriculture and Consumer
Services, we have been able to pursue various aspects
of this study during the past 30 years. For this oppor-
tunity, and for their encouragement and understand-
ing, we thank the following former and present admin-
istrators of this organization: The Honorable Doyle E.
Conner, Commissioner of Agriculture; Dr. W. G.
Cowperthwaite (deceased), H. L. Jones, Dr. S. A.
Alfieri, and R. Gaskalla, Directors, Division of Plant
Industry; and H. A. Denmark, Chief of the Bureau of
Entomology.
We also have had close cooperation from our past
and present colleagues in the Bureau of Entomology:
G. W. Dekle, Dr. G. B. Edwards, Dr. E. E. Grissell, Dr.
A. B. Hamon, Dr. J. B. Heppner, Dr. F. W. Mead, Dr.
L. A. Stange, Dr. M. C. Thomas, and Dr. H. V. Weems,
Jr. They provided specimens, advice, and companion-

ship on many field trips.
In addition to the support from the Florida Depart-
ment of Agriculture, organizational support and grants
were received by the senior author from: 1) The
National Science Foundation (for a summer trainee-
ship at the Organization for Tropical Studies in Costa
Rica; for participation in the Summer Institute for
Systematics at the Smithsonian Institution; for 2 months
of study at the Illinois Natural History Survey; and for
a week of study at Texas A. & M. University). 2) The
Florida Game and Freshwater Fish Commission and 3)
the U. S. Army Corps of Engineers for participation in
surveys of the Cross Florida Barge Canal area and the
Osceola National Forest. 4) The University of Florida,
Department of Entomology & Nematology, and the
Institute of Food & Agricultural Sciences, for the
opportunity to use the scanning electron microscope
and for graduate student assistance over the past sev-
eral years. 5) The Illinois Natural History Survey for
the opportunity to spend 2 months studying there, and
continued support. 6) The Texas A. & M. University at
College Station (Dr. H. R. Burke and E. G. Riley) and
at Dallas (Dr. R. L. Crocker and Dr. J. E. Reinert) for
the opportunity to study for a week at each collection.
We are indebted to most of the major museums in
the United States for loans of specimens or use of their
facilities during personal visits. We apologize for any
inadvertent omission in the following list (abbrevia-
tions are those used in the text and proposed by Arnett,
et al., 1986); names of curator(s) who provided assis-
tance are listed after the abbreviations: American
Museum of Natural History (AMNH), M. A. Cazier,
M. Statham, P. Vaurie, L. H. Herman; Archbold Bio-
logical Station (ABSC), M. A. Deyrup, J. N. Layne;
Canadian National Collection (CNCC),E. G. Monroe,
H. A. Howden, E. C. Becker; Chicago Field Museum
Natural History (FMNH), R. L. Wenzel, H. S. Dybas;
Florida State Collection of Arthropods (FSCA), H. V.
Weems, Jr.; Illinois Natural History Survey (INHS),
W. E. LaBerge, J. K. Bouseman; Museum of Com-
parative Zoology, Harvard University (MCZC), P. J.
Darlington, S. Shaw, J. F. Lawrence; Ohio State Uni-
versity (OSUC), J. N. Knull, F. J. Moore, C. A.
Triplehor; Purdue University (PUIC), Leland Chan-
dler, R. H. Arnett, Jr.; Texas A. & M. University
(TAMU), H. R. Burke, E. G. Riley, R. L. Crocker;
United States National Museum (USNM), O. L.
Cartwright, R. D. Gordon; University of Florida (in
FSCA), T. J. Walker, J. E. Lloyd; University of Michi-
gan Museum of Zoology (UMMZ), Barry O'Conner,
M. F. O'Brien, T. H. Hubbell, R. D. Alexander.
We are also indebted to the following individuals

for various assistance, including the loan or donation
of specimens from their private collections: R. H.
Arnett, Jr., R. M. Baranowski, Bernard Benesh, L. J.
Bottimer, O. L. Cartwright, Neil Chernoff, P. M.
Choate, Jr., L. R. Davis., Jr., M. A. Deyrup, N. M.
Downie, B. K. Dozier, H. L. Dozier, Jr., M. Drucken-
brod, T. Fincher, J. H. Frank, C. A. Frost, S. W. Frost,
E. J. Gerberg, R. D. Gordon, D. H. Habeck, G. Halffter,
A. R. Hardy, E. I. Hazard, C. Hilfiker, H. E. Hinton, H.
F. Howden, O. E. Hunt, R. L. Jacques, P. E. Landolt,
J. W. McReynolds, A. Martinez, C. W. Mills, III, F. J.
Moore, C. W. O'Brien, D. R. Paulson, S. B. Peck, P.
Reyes C., E. G. Riley, P. O. Ritcher, H. H. Samol, M.
W. Sanderson. Joe Schuh, P. E. Skelley, B. J. Smittle,
W. R. Suter, M. C. Thomas, P. A. Thomas, D. W.
Thornton, R. H. Turnbow, K. W. Vick, W. W. Warner,
F. N. Young, Jr.
Light traps were a primary source of specimens
and many individuals cooperated by operating traps
and preserving samples during the past 30 years and,
we sincerely thank them: T. R. Adkins, W. W. Baker,
R. M. Baranowski, W. L. Beers, E. N. Bishop, D. C.
Blanton, F. S. Blanton, A. H. Boike, T. W. Boyd, P. E.
Briggs, R. E. Brown, S. H. Brown, F. A. Buchanan, E.
M. Collins, Jr., H. W. Collins,L. Collins,E. E. Crooks,
G. W. Desin, C. F. Dowling, Jr., P. C. Drummond, G.
B. Fairchild, H. M. Faircloth, E. H. Frederic, Jr., A. E.
Graham, V. K. Gupta, J. C. Hanlon, D. L. Harris, J.
Hayward, E. I. Hazard, L. A. Hetrick, E. W. Holder,
Jr., N. Holler, E. G. Kelsheimer, R. L. King, J. H.
Knowles, M. Lutrick, R. T. McMillan, D. L. Mays, E.
S. Mercer, E. P. Merkel, M. L. Messec, T. Morris, A.
L. O'Berry (Fields), J. W. Patton, J. W. Perry, A. M.
Phillips, W. H. Pierce, J. E. Porter, W. C. Rhoades, B.
J. Smittle, R. W. Swanson, W. B. Tappan, L. W.
Taylor,W. H. Whitcomb, J. R. Wiley, J. W. Wilson, D.
P. Wojcik, D. O. Wolfenbarger.
The two localities which produced the most rec-
ords are Gainesville and Tall Timbers Research Sta-
tion (Leon Co.), and we especially thank the present
and past staff there for many favors: E. V. and Roy
Komarek, W. W. Baker, D. L. Harris, L. Collins, W. H.
Whitcomb, Awinash Bhatkar, and Mary Arnett.
We thank the many Division of Plant Industry
(DPI) and U. S. Department of Agriculture inspectors
in Florida who provided assistance and specimens.
Although space does not permit listing all of them
here, most are listed as collectors in the Appendices.
We thank many other DPI employees for their contri-
butions: for photographic assistance, Jeffrey Lotz; for
art work, John Corkery; for editorial assistance, K. R.
Langdon, F. W. Mead, M. C. Thomas, H. V. Weems,

Jr., N. El-Gholl, J. O'Bannon; for library assistance,
Irene Ayres, M. Batcy, C. Edwards, Louise Henley,
June Jacobson, A. Kolesar, Alice Sanders; for secretar-
ialwork, I. Ayers, E. Manning, D. Proveaux, J. Temple,
F. Williams, S. Yocom; for microscope slide prepara-
tion, A. L. Fields and E. Ostanik.
For assistance with the scanning electron micro-
scope (SEM), we especially thank: ThelmaC. Carlysle
(USDA, retired) for many volunteer hours and techni-
cal aid; the SEM supervisors: Dr. G. Erdos and Donna
Williams, Botany Department, University of Florida;
and especially to entomology graduate student P. E.
Skelley for his mastery of both the critical point drier
preparation technique and his adept use of the SEM
itself.
During 2 months of study at the Illinois Natural
History Survey, the following individuals contributed

in many ways: J. K. Bouseman, A. R. Brigham, A.
Eckhoff, B. Gillics,G. Godfrey, C. Heister, S. Heydon,
A. Kirts, K. McGiffen, S. L. Passoa, R. L. Selander, J.
Sherrod, D. & S. Voegtlin, and M. L. Williamson.
Although mentioned in other capacities, we must
separately express special thanks to 4 individuals: Dr.
Milton W. Sanderson to whom we dedicate this vol-
ume, for his undaunted support in every possible way,
and for his many years of devotion to the study of this
genus; Paul E. Skelley to whom we dedicate one of the
new species, for his assistance in every aspect of the
study; Dr. Michael C. Thomas for his computer exper-
tise and personal sacrifices to facilitate the camera-
ready copy; and to Frances Williams for her meticu-
lous conversion of a handwritten manuscript to an
accurate computer version; and to them all for their
valued personal friendship.

FRACT

In this faunal study, data are presented for 54 species of Phyllophaga recorded from Florida. Two new
species, pseudofloridana and skelleyi, are described, and 9 other species are recorded from the State for the first
time: anxia LeConte, apicata Reinhard.forbesi Glasgowfoxii Davis, implicit Horn,perlonga Davis,profunda
Blanchard, puberula DuVal, and yemasseei Cartwright. In addition, illustrations are provided for the first time
of the female genitalia for 4 species: clemens Horn, elizoria Saylor, georgiana Horn, and okeechobea Robinson.
Two species [P. murrea Sanderson, previously known from a unique female, under P. elongata Linell, and P.
vanalleri Schaeffer, originally described as a subspecies of obsoleta, under obsoleta (Blanchard)] are syn-
onymized.
Scanning electron microscope photographs are provided for the male (4 or 5 views) and female (2 views)
genitalia, as well as other relevant taxonomic characters. Data are presented for each species under the following
headings: synonymy, type locality, diagnosis, description, taxonomic notes, distribution (Florida and United
States), biology and ecology, adult host plants, immatures, specimens examined, and selected references. Maps
showing both Florida and United States distribution are provided for each species.
Keys are provided for the adults of 54 species, and for the known larvae of 23 species. Eight tables provide:
1) an alphabetical checklist of figures and maps; 2) a chronological list of Florida Phyllophaga descriptions; 3)
the arrangement by "groups" of various authors; 4) a summary chart of life cycle data; 5) an alphabetical list and
references to known larvae of Florida species; 6) a list of 17 misspelled names of Florida species, and their
citations: 7) an annotated alphabetical checklist of the 96 names used for Florida species; and 8) a summary of
22 basic taxonomic character states. Specimen label data are listed in the Appendices (1-39) as is a list of the
figures (40), and the Bibliography contains 376 references.

PLATE' FIG 1-12 PHYLLOPHIAA MALE GENITALIA (CAUDAL), Une-0.Smm
.mE
,'. ,. ..
/i/il

PLATE 2. FIG 13-24 PHYLLOPHAGA MAE GENITALIA (CAUDA)U. U-ne5.mm

HI

HI
IB
1 1--4. r
r1 1-4S u,

idI
11a

PLATE6. FIG 6172,

(VENTRAL), ULn0.5mm
II
II
M I .
II

FIG.73-8 PHYLLOPHAGA MALE GENITALIA (VENTAL) b OSmm

I.
I--I
HUMMl

Em3

U

AGA MALE GENITALIA (VENTRAL), LneO5 mm

El
Is oa/l

PATE10: FIG. E 09-120. PHYLLOP AGA MAE GENITAA (VENTRAL), LI NeNSmm

mi,

.

:FI 121-132. P HAGA MALE GENITALIA (DRSAL) Un mm

--H ul.ebala
I H^I11

PULTE12: FIG.133-144. PYLLOPHAGA MALE GENWmIA (DRSAL), U .mm

15

Il
I

II
//

11

I.I

PTE: FIG.16180. PHLLOPH LE EN LI (RSL), Le 5mm

,--
I.I

iii
I,.
m -- 11 bl~a2 n~a.^a) Iouu(om
1.E
E I

I./

BHH
-- _,fli.eg

: F1G.205.216 PHYLLOPHIGA MALE GENITAL[ (RIGHT LATIERAL},. Une-O.Smm

I.l
1mm

PLATE19, F1G.217-22 PHYLLOPHAG& MALE GENITAIA (RIGHT LA0TERA0L), LAn0 .Smm
I.IE
I.B
i.E
/Jill
..... . ... .

PUTEM FIG 229-U). PHYLLOPHAGA MALE GENITALIA(RIGHT LATEL). Um.0.5mm

PLATE21: FIG.241-252 PHYLLOPHAGA MALE GENITALIA LEFT LATERAL. UL-O.5mm

i.i
I.E
Em.a
//~u--~0rhl ailN

PUTE u: FIG. 263211. PHYUOPH*GI FEM*LE GENIT*LI* (YENTRPL UIERBJ. Un~O.Smm

,rotrl -- w.lad~*~al -- 166.~l~f~vratr~

-- u~.nr~.lrt~re -- ~s.plut,~t~RI

l~0.d.mlognobl --llt.dYPlt~vnUl

il1~I 113. e~m~ar

PUTE 21: F G 215-286 PHYLLOPHIGI FEMALE GENITALIA (VENTRIL LATERAL). Une-o.s mm

~4*'0rm ~1I~.EulYWorn~~.rrotnl ~lll.drblllgrmml

Y4UI~~I ZIP. cu~~llrorn21U~l ~0. ~bU4~U~I

Isrntrl 2Bl.drtll.rrobl

I

IHI
un a

U

I.E
EHu
I,~~ l. Ynn

IG.335 346. PHYLLOPHAGA FEMALE GENITALIA (VENTRAL-LATERAL), Une=0.5mm

.043:FG*I38 HLOHIO E*EONILLAIET

PHYLLOPHAGA FEMALE GENITALIA (VENTRAL- LATERAL), UrlO 5mm

-.-n -n.. __m,.
El

PLATE33: FIG.379390. PHYLLOPHAGA ANTERIOR TARSAL CIWS A ELY

MiW
El

Ii
li

UI

FLATE35 FIG.403-HB. PHTHLOPHAGA MALE ABDOMENS CENTRAL)
BE
.,.1.^ .^.^^^^B

Eu
uMp
mmm^
y '^.^
cmo ESSSS~

aemula
anxia (so.)
anxia (no.)
apicata
bruneri
clemens
clypeata
crenulata
cupuliformis
debilis
diffinis
dispar
drakii
elizoria
elongate
ephilida
floridana
forbesi
forsteri
foxii
fraterna
futilis
georgiana
glaberrima
gracilis
hirticula
hornii
ilicis
implicit
infidelis
knochii
latifrons
Iota
luctuosa
mariana
micans

Table 1. Alphabetical checklist of figures and maps.

Male Genitalia Female Genitalia Text Map
caudal ventral dorsal lateral ventral lateral Figures FL US
1 61 121 181 263 266 464 465
2 62 122 182 264 267 466 467
3 63 123 183 467
4,5 64,65 124,125 184,185 265 268 403,468,470-1 474 475
6 66 126 186 269 272 404, 426-7, 457-61,476-7 478 479
7 67 127 187 270 273 391 480 481
8 68 128 188 271 274 482 483
9 69 129 189 275 278 484 485
10 70 130 190 276 279 486 487
11 71 131 191 277 280 392,418 488 489
12 72 132 192 281 284 453 490 491
13 73 133 193 282 285 379, 393 492 493
14 74 134 194.243 283 286
15 75 135 195 287 290 8, 419s20.462-, 44 597
5,50 5688, 496 497
16 76 136 196 288 291 498 500 501
17 77 137 197 289 292 394,455 502 503
18 78 138 198,244 293 296 504 505
19 79 139 199 294 297 506 507
20 80 140 200, 245 295 298 452 508 509
21 81 141 201,246 299 302 432,510-516 517 518
22 82 142 202,247 300 303
23 83 143 203 301 304 395,405 519 520
24 84 144 204 305 308 380,406 521 522
25 85 145 205 306 309 396,422 523 524
26 86 146 206 307 310 525 526
27 87 147 207,248 311 314 385 527 528
28 88 148 208.249 312 315 419,529 530 531
29 89 149 209,250 313 316 532 533
30 90 150 210,251 317 320 534 535
31 91 151 211,252 318 321 397 536 537
32 92 152 212,253 319 322 538 539 540
33 93 153 213 323 326 407,423,430,433-8,541-2 543 544
34 94 154 214 324 327 381 545 546
35 95 155 215,254 325 328 398,408 547 548
36 96 156 216 329 332 -49 550
37 97 157 217 330 333 382.399.409

murrea
obsoleta
okeechobea
ovalis
panorpa
parvidens
perlonga
postrema
profunda
prununculina
pseudoaloridana
puberula
quercus
schaefferi
skelleyi
subpruinosa
taxodii
tecta
tristis
ulkei
uniforms
yemasseei
young

Table 1. Alphabetical checklist of figures and maps (cont.).

Male Genitalia female Genitalia Text Map
caudal ventral dorsal lateral ventral lateral Figures FL US
331 334 551
38 98 158 218 335 338 383, 410 552 553
39 99 159 219 336 339 389,494,588 554 555
40 100 160 220,255 337 340 416 556 557
41 101 161 221 341 344 413,414,499 558 559
42 102 162 222 342 345 439-441 560 561
43 103 163 223,256 343 346 562 563
44 104 164 224,257 347 350 451 564 565
45 105 165 225,258 348 351 411,566 567 568
46 106 166 226 349 352 386, 400, 446-450 69 570
47 107 167 227.259 353 356 571,572,573 574 575
48 108 168 228 354 357 576 577
49 109 169 229 355 358 401,428-429,431,578 579 580
50 110 170 230,260 359 362 581 582
51 111 171 231 360 363 390.424-5583,585,587-8 589 500
52 112 172 232 361 364 591 592
53 113 173 233 365 368 593 594
54 114 174 234,261 366 369 387 595 596
55 115 175 235 367 370 421,454,469,4723 597 598
56 116 176 236,262 371 374 417. 442-445 599 600
57 117 177 237 372 375 402,415, 601,602 603 604
58 118 178 238 373 376 605 606
59,60 119,120 179,180 239,240 377 378 384,456,607 608 609

The figures for theplates wereproduced with a camera attachmenton the scanning electron
microscope. The male genitalia are all shown in 4 views, and the asymmetrical ones in 5 views
(right and left lateral). The female genitalia are all symmetrical and are shown in 2 views (ventral
and right lateral). Genitalia of drakii,fraterna, and micansare illustrated for comparison, although
not recorded from Florida. Label data for the illustrated specimens are provided in Appendix 39.

INTRODUCTION

Probably no other genus of North Aarnicas in-
se. s so universally reedgmed as ame the "May or
JuneBugs". Folk sngsandna y rhymea have used
themas a themc, and chldie play with them Itis.no
surpng. smne adult beetles ofn sw.7 by the

desetlve larvalfI Fewlaymen ralz that the

Imwaeoften spend 2 m4 years beneath thesoil, feeding
on marsof variousplants, espeeially h, sad uW fo

Theyal "do notrealise, hatthere isnoton e "Jam
Bog but the re re
Undecd Smas and 54 FZ= = ,,ve I. the fro
Flonds, Bsaadcfly, Phyllophag am allquite similar i
Ms. "Id "" "'. '~

INN,~I II L I Mrll. L

1.s hN, OII~ofIINS(May bN .) p IN
The name hyllophaga comes firm the Gr-kr Phyll
led, and phaga = n, min reference to teir sriou
defolation of ornour hardwood mmst in the aist and
midwest,
Because of their general similarity and grant
arandusee, they no, -gleetdby suirycoll-,to and
sm, thought Mo be difficult to ientif by many pmf,,.
nimial gnIINIINoids However, large, wallIdevel-

plater 1.32). The genus provides an arealleml oppor-

eumo5, 1ning beh5iun,, fts5

Forl at of Presentatima

The general format is that used for Part I of the
"Scaab Beefles oFlorid," (Woodialf, 1973). Same
noodificabors emad ssthipsvolume dNals with
a single ganus. Nsrese liability hasb..flared 1o
genitalia ithemminaios rthe, than trditional krey,, I-
though key. In- idd far both adults and larvic.
The genitalia f' gars plaNed st the beginning e-
cmIse N fIi i, inaprna and to enable their number-
.1; before woplebom of the -
This volume was mac o opue ndsb

re -..-a U~ nand ricintimi It se eliminate

Beetimic of inare reponsitnllmr, of the co-
authors, and in avoid confusion and passive voice, the
poonoun t is used in ... f are tart. There a.P few
saaabo's (e.g., ca lta of type spmveron in
other musamms) when only the senior author was

S1~

involved; in thew assisthepronoun I ,. sM. The
spezimmn dama were produced Beart and formattd lawo
the Appendwe ,. as hckg, .. d for the test discm-
sions. TheBibllogrphy was completedrest and the
peitinum, of-., .e were cited by ilaxles n he Sa-
lected References' Th. individual spa~ihr Lremucma

computrmftcbh~p,,,, TheDivibucamMapl
were than cmpleted and ost fiareo dded. The
Intrudoc' - d India er written his,

adaptations of anstatg koes, odiftled for the Morid,
feums. with addimm. and deletons as required.
Aflthmigh the adult beetle, have win, awcelleal
flIa l chmantes. many bfige yycannot be identified
with ase or certainty without reference L. the gamam

caution for saverl ressions: 1) am thaln half the
1mvw mfolorid. spaci. are ..know.; 2) most thame--
tons wre based on few hearing, and variation has att
studied; and 3) -a species (ag., ..W.) an,

the makirs.
11 ustratism,: Most of the half-tons figunce were,
anwe on the starting clecrom afirmstope (dinsil,
.ad. Madmi& & Materias). Lin. me bellbol

original and copies from Al.i. g literature (par.cu
Italy larval characters). Th, scancfor Lbla im, are
always indicamd. Photographsofminingbehavio
were night in the field by JefL and R. E.
Wominuff.
Sp~eci Aounts: Each~Aa Flord species (asp
these irested in pumammnble ard Potential Records)
was treaed inthefollowing forasa
i) Synonymy: All previous name combinations
i, liAd in ch.g..A.gica. order,.each army followed
by the antho, date. and page (complete rinationsA s
found inhBibliography). A comma belweenL Al
..me and the muthe, midicides dam be -as responsible
for the timecombination, bar,, no the uthorrifta

A2) Type loAlity A When A iwasble, to d
migincaactliseildies 'they are listed m quo as inthe
rgiab description. In somecases, no specifilaId-
ity was give, (e g., uro and the presumed Recality

3) Diagnosis: bffies werb made Ml maLk Has

flllcly (esternlly) lami. l which appear to be
osly laled by glel i al hummons. Ia sim.. ilar
spcisme liAA,,andAreferenceisAmadeItothe
rent figimc. Oc-imialply supplemental mmrphalogi-
.I ct -flesnr mddd,

over 50 similar species would rarely be helpful and
would occupy an inordinate amount of space. We
believe this is a special group in which the saying, "a
picture [of the genitalia] is worth a thousand words",
aptly applies. We have discussed the following char-
acters for each species: Length was measured from the
tip of clypeus to the tip of elytra (not abdomen, which
may move, especially after dissection). These num-
bers represent the minimum and maximum length in
our series of specimens to the tenth of a millimeter.
Width was measured at the maximum point (variable
because some specimens have elytra slightly separated
at the suture and therefore appear wider). These
numbers represent the minimum and maximum in our
series of specimens. Numbers in parentheses, fol-
lowed by an author's name, are extensions (either
direction) of our measurements and the reference for
them. Shape: Thegeneral form isdescribedasoblong,
oval, parallel, widest behind, convex, or subdepressed.
Color: The species of Phyllophaga vary from pale
yellow through various shades of brown to nearly
black. Unfortunately no standards have been estab-
lished for these colors, and they are difficult to express
precisely. The older, latinized terminology, used by
LeConte, Hor, et al., has been followed by most
workers. Even though we don't believe the terms
convey the shades of color effectively, we have no
better system at present. However, many species are
distinctively colored and often can be rough sorted
from mixed collections by color alone. Vestiture: The
surface coating is either pubescent (hairy), pruinose
(velvety), or glabrous (smooth). Occasionally a spe-
cies (e.g., aemula) has the surface pruinose and pubes-
cent. The hairs setaee) of pubescent species may be
recumbent or erect, short or long, dense or scattered,
and may differ on head, pronotum, elytra, and abdo-
men. Glabrous species are shiny or dull. Antenna:
The most obvious character is the number of segments
(from 9 to 10), and the sexual dimorphism in length of
the club (larger in males). The length of the male club
is given in relation to the previous segments of the
scape (stem). Clypeus: The front margin is either
entire (not indented at middle) to emarginate (with
median indentation deep, shallow, or moderate). The
margin is reflexed or not, the edge raised prominently,
moderately, or slightly. Tarsal claws: The claws are
curved slightly or strongly. They are cleft in the
subgenus Phytalus (georgiana & obsoleta), but toothed
in nearly all others. The tooth position is most fre-
quently median, but may be basal, nearer the base
(subbasal), or nearer the tip (antemedian). The tooth
shape may be acute, blunt, and prominent or reduced.

It appears to be absent in males of panorpa. Male
posterior tibial spurs: Normally there are 2 enlarged
terminal spurs, the lower (outer) one may be fixed
(unarticulated) (fig. 391, 392, 394, 400,402) or mov-
able (fig. 393,401) (articulated), but the upper (inner)
spur always is movable, normally longer. The lower
spur occasionally is modified into a twisted shape (fig.
395) or reduced to aborted (fig. 396, 500,402). The
shape of each spur is described as are their relative
lengths to each other and to the first tarsal segment.
Genitalia: Because there is no standard terminology
and there are several basic genitalia types, no lengthy
descriptions are attempted. However, figure refer-
ences for the various views are cited, and these should
provide positive means for identification ofboth males
and females.
5) Taxonomic Notes: Any information was in-
cluded here on synonymy, type specimens, questions
about validity, variability, etc.
6) Distribution: We provide maps, and reference
to them, for both Florida and the U.S. We have divided
them so that discussions on records are clearer. Florida
records are based on specimens examined only, unless
a specific questionable literature record is discussed.
All specimen data are either listed under SPECIMENS
EXAMINED (10 or less records) or in the Appendices
(10 or more records). The U. S. distribution was
basically plotted from the literature, but trying to
locate specific records in order to place dots accu-
rately. Questionable or unusual records are shown on
the map by a question mark.
7) Biology & Ecology: General remarks are given
first on such things as seasonal activity, abundance,
and other notes. These are followed by 2 subdivisions:
a) Adult Host Plants: These are primarily literature
records, normally with Luginbill & Painter's (1953)
list by family first. It is then followed by other host
records, usually by state, with author and date citation.
Few adult hosts were recorded in our study, partly
because of the apparent non-specificity of most spe-
cies and the ease by which they may be collected at
light; b) Immatures: The known larvae are described
from the literature. Little time or effort was spent on
rearing, so there is little original data here. However,
the pertinent literature is cited to aid the reader in
obtaining information.
8) Specimens Examined: For those species with
less than 10 records, the data are listed here; the others
are only summarized, with the total number and refer-
ence to the full data in the appropriate Appendix.
9) Selected References: Except for catalogues or
checklists, we have attempted to cite all references to

Table 2.
Chronological List of Florida Phyllophaga Descriptions
(only valid species are listed)

1781. tristis Fabricius
1792. crenulata Froelich
1801. quercus Knoch
1801. ilicis Knoch
1801. hirticula Knoch
1817. knochii Shoenherr & Gyllenhal
1825. ephilida Say
1850. anxia LeConte
1850. futilis LeConte
1850. profunda Blanchard
1850. uniforms Blanchard
1850. glaberrima Blanchard
1850. diffinis Blanchard
1850. obsoleta Blanchard
1855. forsteri Burmeister
1855. prununculina Burmeister
1855. gracilis Burmeister
1855. dispar Burmeister
1856. latifrons LeConte
1856. puberula DuVal
1856. parvidens LeConte
1856. debilis LeConte
1884. subpruinosa Casey
1885. georgiana Horn
1887. clypeata Horn
1887. clemens Hom
1887. postrema Horn

each species by author and pages (with figures, tables,
plates, and maps). Full citations appear in the terminal
bibliography. We undoubtedly have missed some
references, but we believe the section is relatively
complete.

Historical Resume

The first list of Florida Phyllophaga was that
included in the "Coleoptera of Florida" (Schwarz,
1878). Six described species (latifrons, cerasina,
glaberrima, micansfraterna, and tristis), and 4 unde-
scribed species were recorded under the generic name
Lachnosterna. Dozier (1918) listed 5 described spe-

1887. infidelis Horn
1887. luctuosa Hom
1887. implicita Horn
1887. aemula Hom
1889. ulkei Smith
1889. hornii Smith
1896. elongata Linell
1916. forbesi Glasgow
1920. perlonga Davis
1920. foxii Davis
1924. taxodii Langston
1924. cupuliformis Langston
1928. lota Luginbill
1929. mariana Fall
1932. bruneri Chapin
1935. young Cartwright
1937. elizoria Saylor
1937. schaefferi Saylor
1938. floridana Robinson
1939. apicata Reinhard
1939. ovals Cartwright
1944. tecta Cartwright
1944. yemasseei Cartwright
1948. okeechobea Robinson
1950. panorpa Sanderson
1989. pseudonloridana Woodruff & Beck
1989. skelleyi Woodruff & Beck

cies (prununculina, glaberrima, parvidens, latifrons,
and micans) and 1 undescribed species "near nova"
from Gainesville.
The first extensive list (29 species) was produced
by Blatchley (1929) in his "Scarabaeidae of Florida".
Young and Thames (1949) reported 41 species in their
"PreliminarylistofthePhyllophagaofFlorida". The
monograph on the 'May Beetles of the United States
and Canada" (Luginbill & Painter, 1953) contained
records of40 Florida species. In Part I of the "Scarab
Beetles of Florida", Woodruff (1973) listed 40 de-
scribed and 2 undescribed species of Phyllophaga.
In the present study we recorded 54 species,

including 2 new species, 9 new State records, and 2
species placed in synonymy; 16 additional species are
listed as questionable or potential.

Methods and Materials

Over 100,000 specimens of the genus Phylloph-
aga were examined for this study. Because the geni-
talia are the most reliable taxonomic characters, these
were normally dissected. Since many species are
extremely common, voucher specimens of these often
were preserved in alcohol (70% isopropyl) rather than
conventional pinning.
Genitalia: Dissection of genitalia from these large
beetles is an easy operation. The last ventral segment
and the pygidium are pried apart so that the genital
opening is accessible. Generally forceps are an ade-
quate tool to extract the genitalia on fresh specimens.
However, an insect pin or dissecting needle is handier
for females and alcohol preserved specimens. Dried
specimens must be relaxed in a humidity chamber, in
an ultrasonic machine, or in boiling water, prior to
dissection.
On specimens to be retained in alcohol, the geni-
talia were exposed sufficiently for identification, but
were left attached to the internal membranes. For
pinned specimens, the genitalia may be handled the
same way, but they often are mounted on traditional
triangular card points or preserved in microvials with
glycerine. Some minor cleaning of the male claspers
and the female pubic process may be required to see all
the features, but clearing with potassium hydroxide is
not required.
The male internal sac or aedeagus usually is
hidden within the basal piece and often requires everting
in order to study it. In some species (e.g., apicata,
bruneri, tristis, youngi, the sac is always somewhat
everted and normally visible without further dissec-
tion (fig. 4,59,64,119,124,179,184,239). Presently
most taxonomic characters are found on the para-
meres, but limited study suggests that excellent char-
acters may be found on the aedeagus. These may be
useful in clarifying some of the similar species, such as
the fraterna complex.
During our early studies we made microscope
slide preparations of these, but their 3-dimensional

*Mention of a proprietary name does not necessarily
imply endorsement of the product or company by the State
of Florida.

shape was lost, and relative positions of characters was
obscured. In order to study them "in the round"
techniques were developed (by P. E. Skelley) to keep
the soft parts rigid. The sacs were everted either by
dissection with jeweler's forceps and hook-tipped
minutes (for previously dry specimens that had been
relaxed in detergent water), or by squeezing the sur-
rounding muscles to inflate the aedeagus by osmotic
pressure (for freshly killed specimens). The inflation,
especially on the relaxed specimens, could be in-
creased by injection of 70% isopropyl alcohol through
a hypodermic syringe.
Once inflated, they were dehydrated through several
changes of ethanol (80-90-100-100%). Dehydration
required at least 10 changes for fresh specimens (from
10-100%). They then were dried in a critical point
drier (Tousimis, Samdri 780A*), so that all parts
remained rigid and in place. They were mounted
finally on card points which were glued to a stub for
eventual study and photography under the scanning
electron microscope. Our studies are very preliminary
for the aedeagus, and we only illustrate a few to show
how significant they may be and to encourage their
future study (fig. 433-450, 583, 601, 602).
Illustrations: The genitalia are at a size range and
complexity that conventional photography is difficult
(particularly depth of field). No single view of the
male genitalia is adequate to clarify characters in most
species, and line drawings of several views each (about
380 required) were planned. However, the use of a
scanning electron microscope enabled us to prepare
much better and more detailed illustrations.
Specimens (genitalia) were cleaned in an ultra-
sonic cleaner, glued to an aluminum stub, and coated
with 24 karat gold (fig. 412) (approximately 150
angstroms thick) in an Eiko Engineering IB-2 Sputter
Coater*. They were placed then in the chamber of the
scanning electron [SEM] microscope (Hitachi S-450*)
with a camera attachment. Most study was done at 20
kilovolts, permitting a minimum magnification of 16
times. After specimens were oriented, they were
photographed on Polaroid (55 Professional, positive/
negative*) 4x5in. instant sheet film. Negatives were
developed and stored with the positive print for later
darkroom work. Scale lines (in microns) were pro-
duced automatically by the microscope and appeared
on the photographs. Over 1000 photographs were
taken, in order to obtain the final 548.
All final prints for the plates (1-35) were made
from the polaroid negatives, enlarged or reduced for
uniform size, and improved for reproduction, with
standard techniques in the darkroom (for which we are

greatlyindebted to JeffLotz). They then were cropped
to a uniform 2 1/4in. square for composition into the
plates.
Because the backgrounds of the SEMphotographs
were not uniform, obscuring certain features, we de-
veloped a technique for painting them black (see
beforeandafter, fig.413,414). The finishedproductis
therefore a combination of photography and artwork.
Painting was done on the photograph surface with fine
camel's hair brushes (0000 to 2), using the opaque
black water soluble paint (No. GY9) of a Gamma
Retouch Set (25/8, M. Grumbacher, Inc*.). Dirt and
damaged areas also were retouched with the other 7
shades of white and gray in this set. Over200 hours of
painting time were required.
The finished illustrations were mounted on heavy
stock in the format of the plates, using strips of double-
facedcarpet tape (Super Stick,Superior, Union, Mo*.).
Legends for the plates, scale lines, and the squares for
the illustrations were generated on a computer with
Aldus Pagemaker*. The scale lines were measured to
the nearest millimeter on the polaroid print and then
drawn to that length on the computer. The sheets with
figure legend, scale lines, and outlines of the squares
were used as an overlay negative by the printer to crop
the photographs uniformly, when combined with the

half-tone negatives. Windows, the size of the finished
half-tone areas, were made of ruby lith to match the
computer generated squares. Plates 1 to 34 were
finally reduced by 6%, but Plate 35 was reproduced at
original size.
Halftone negatives of 150 line screen, using the
Duo-tone process, were prepared by Storter Printing
Co., Gainesville; printing was done by Paramount-
Miller Graphics, Jacksonville, Florida.

Collecting Techniques

Perhaps this section might appear superfluous,
since even children can easily collect "June Bugs"
around the lights at night. Probably all the Florida
species are attracted to lights, if they are within a
reasonable distance of their habitat. Morethan95% of
our specimens were collected with blacklight (ultra-
violet) light traps of various design (see U. S. Depart-
ment of Agriculture, 1961).
However, many of the rare species either have
restricted habitats, or we know too little about them.
Some (e.g., taxodii on cypress) are so host specific that
special effort must be made to operate traps near the
host. In other cases (e.g., skelleyi in turkey oak scrub)
the general habitat must be located to collect speci-
mens.
Because they feed and mate at night (with the
possible exception of okeechobea), they are collected
easily from their hosts, with the aid of a headlight
(flashlights require a hand, and both are needed to hold
the container and to collect specimens). Beetles usu-
ally are so intent on feeding or mating that they are not
disturbed easily, and they may be picked like grapes.
Often more diversity and specimens may be collected
from foliage (in a comparable time period) than a light
trap in the same area. Usually we try to use both
techniques.
Specimens may be collected in killing jars with
cyanide, ethyl acetate, or lighter fluid, or they may be
preserved in alcohol (70% isopropyl preferred). Be-
cause they are large and exude body fluids on death,
the latter technique is better, providing adequate pro-
tection until genitalia can be dissected and specimens
can be mounted.

GENERAL ACCOUNT
OF THE GENUS PHYLLOPHAGA

This is one of the largest genera of animals in the
ig. 415. Phyllophaga uniforms: adult beetles feeling on elm UnitedStates,whereover200speciesoccur. Anequal
foliage at night.

number may be found in Mexico and Central America.
Because there are so many species, generalities are
difficult to make. They occupy many ecological
niches, andevery major habitat(exceptaquatic) seems
to have been invaded. Since they are often economic
pests in both adult and larval stages, it is imperative to
know which of the many species is involved, so that
control strategies can be planned effectively.
Taxonomy: The United States fauna is fairly
well-known, partly due to the work of M. W. Sander-
son (1937-1958, and unpublished) and the faunal study
byP.LuginbillandR. H. Painter (1953). This is not the
case in other areas (e.g., Mexico and the West Indies)
where up to 50% of the species may be undescribed.
The Florida fauna was well-known previously, and the
2 new species described herein are the first in nearly 40
years.
However, the U. S. fauna was not always on such
a firm foundation. Historically, our species were
placed in numerous genera from 1781 until 1916 and
namedby 25authors. They are shown in chronological
order in Table 2. The name of the genus is firmly
established now, but past confusion about its validity
contributed to some homonomy and disagreement
among various workers. Glasgow (1916) provided a
detailed review of the confusion and was instrumental
in stablizing the name.
Briefly, the problems revolved around Harris'
(1827) original description of Phyllophaga in a foot-
note, which contained several valid species, but was
not accompanied by a description. The International
Rules of Nomenclature clearly recognize this as an
"indication", and the name should be recognized for
these insects. The type species, hirticula Knoch, was
chosen by Glasgow (1916).
The date of Harris' description has been listed
often as 1826 (e.g., Glasgow, 1916; Saylor, 1942). In
a later edition of the same book (Harris, 1863:30) he
again used a footnote to cite the original description as
follows: "A genus proposed by me in 1826. It
signifies leaf-eater. Dejean subsequently called this
genus Ancylonycha." However, in 1869 Scudder
published the "Entomological correspondence of
Thaddeus William Harris, M. D." and included a
complete list of his publications, giving the date of the
relevant paper as 1827. The reference was published
in the Massachusetts Agricultural Journal (Reposi-
tory), Volume 10, No. 1, on which the date July, 1827
appears. Harris may have referred to his 1826 date, on
the basis of letters (see Scudder, 1869:17) in that year
or when he submitted the manuscript, but the actual
date of publication should be 1827. However, that date

is of no apparent consequence in the synonymy.
Several prominent early workers (e.g., LeConte,
1856; Horn, 1887; Smith, 1889) believed that Phyllo-
phaga was not valid, because it was not accompanied
by a description. They used the generic name Lach-
nosterna Hope (1837), rejecting both Ancylonycha
Dejean (1833) andPhyllophaga on the same grounds.
In addition to Lachnosterna, LeConte (1856) also
recognized 3 new genera (Eugastra, Endrosa, and
Gynnis) and the Listrochelus of Blanchard (1850)
within what we now call Phyllophaga.
When Horn (1887b) published his monograph of
Lachnosterna, he suppressed LeConte's genera, as
wells Trichestesand Tostegoptera Blanchard (1850).
Saylor (1942) treated the genera related to Phylloph-
aga, establishing the following as subgenera: Phytalus
Erichson (1847), Listrochelus Blanchard (1850), Ch-
laenobia Blanchard (1850), Tostegoptera Blanchard
(1850), Eugastra LeConte (1856), Chirodines Bates
(1888). He also created 3 new genera: Triodonyx,
Clemora,andCnemarachis. Saylorlater(1937)raised
Chlaenobia to generic status, at the same time uniting
the Old World Brahmina Blanchard (1850) and Holot-
richia Hope (1837) with Phyllophaga.
Sanderson (in several papers and pers. com.) pre-
ferred to treat most of the "genera" as subgenera,
breaking them down into "species groups". This
avoids creation of homonomy and keeps the near
relatives together. There appear to be no sets of
exclusive characters which will suffice to segregate
the Phyllophaga relatives into clear cut genera (even
from different continents). Luginbill and Painter (1953)
avoided the problem in their treatment of the U. S.
species by not mentioning subgenera, eliminating the
species of the subgenera Listrochelus and Phytalus,
but including those in the subgenera Tostegoptera and
Eugastra (without reference to these names).
Moron (1986), treating the Mexican species, rec-
ognized the subgenera Phyllophaga, Phytalus, Listro-
chelus, Chlaenobia, Tostegoptera, Eugastra, and Tri-
odonyx, as well as species groups and "complejos"
within the groups. Thearrangements of Horn (1887b),
Boving (1942), Sanderson (mss.), and Moron (1968)
are summarized for the Florida species in Table 3.
The most important breakthrough in the taxon-
omy of this genus came with the discovery that the
genitalia of both sexes offered excellent characters to
distinguishotherwisevery similar species. These were
first used by Smith (1889b), when he illustrated the
genitalia as a supplement to Horn's (1887b) mono-
graph. Prior to that, some of the leading workers had
such difficulties in distinguishing species that they

Table 3.
Arrangement of Florida Phyllophaga according to groupings by various authors.

Horn Boving Sanderson Moron
(1887) (1942) (ms.) (1986)

aemula (Horn)
anxia LeConte
apicata Reinhard
bruneri Chapin
clemens (Horn)
clypeata (Horn)
crenulata (Froelich)
cupuliformis Langston
debilis (LeConte)
diffnis (Blanchard)
dispar (Burmeister)
elizoria Saylor
elongata (Linell)
ephilida (Say)
floridana Robinson
forbesi Glasgow
forsteri (Burmeister)
foxii Davis
futills (LeConte)
georgiana Horn
glaberrima (Blanchard)
gracilis (Burmeister)
hirticula (Knoch)
hornii (Smith)
ilicis (Knoch)
implicita (Horn)
Infidells (Horn)
knochil (Schoenherr & Gyllenhal)
latifrons (LeConte)
!ota Luginbill
luctuosa (Horn)
mariana Fall
obsoleta (Blanchard)
okeechobea Robinson
ovalls Cartwright
panorpa Sanderson
parvidens (LeConte)
perlonga Davis
postrema (Horn)
profunda (Blanchard)
prununculina (Burmeister)
pseudofloridanas n. sp.
puberula (DuVal)
quercus (Knoch)
schaefferi Saylor
skelleyl n. sp.
subpruinosa (Casey)
taxodil Langston
tecta Cartwright
tristis (Fabricius)
ulkel (Smith)
uniforms (Blanchard)
yemasseei Cartwright
young Cartwright

XII
IX

V

XII
(IX)
VI
IX
XV

IV
(IX)

IX
(IX)
VII
Phytalus
IV
VII
XI
IX
XI
X
IX
IX
IV
V
IX
(XII)

(XII)

XII
IX
VII
IX
IV
(IX)

XV

(XII)
IX
VI
(IX)
XVI
IX

1
16
(4)

1
14
9

12

11

11
21a
(21b)
10

11
13
21
21
21
17

21
12

19

1

21
11

2

(14)

4

crenulata
rugosa
setidorsis
Cnemarachis (subgenus)
crassissima
quercus
crenulata
crassissima
crassissima
rugosa
quercus
crenulata
elongata
crassissima
rugosa
crassissima
rugosa
rugosa
crassissima
georgiana
crassissima
crassissima
rugosa
rugosa
rugosa
implicita
rugosa
rugosa
crassissima
crassissima
rugosa
crenulata
obsoleta
crenulata
rugosa
elongata
crenulata
rugosa
rugosa
rugosa
crassissima
crenulata
Cnemarachis (subgenus)
quercus
rugosa
crenulata
crassissima
crassissima
rugosa
setitdorsis
rugosa
crassissima
(Phytalus)
Cnemarachis (subgenus)

anodentata

ephilida

ephilida

Phytalus (subg.)

anodentata

() = placed in Horn's group by subsequent authors.

described the opposite sexes as different species, placed
the same species in 2 genera under different specific
names, and generally added to the problem ofcorrectly
identifying the many species. Horn (1887b) stated that
"Lachnosterha is certainly one of the most difficult
genera in our fauna...," and Riley (1891) published a
paper entitled "On the difficulty of dealing with
Lachnosterna."
When Glasgow revived the name Phyllophaga in
1916, he also was able to establish the synonymy of
many species by reference to genitalia of most type
specimens available at that time. This provided the
nomenclatural stability which has greatly aided later
workers, but unfortunately Glasgow never published
his proposed monograph, nor illustrations of the geni-
talia as he planned.

Generic Synonymy

Phyllophaga Harris

Phyllophaga Harris, 1827:7 (footnote); type: Melolontha
hinicula Knoch, designated by Glasgow (1916).
Ancylonycha Dejean, 1833:160 (in part); type:
Melolontha serrata Fabricius, designated by
Duponchel (1849).
Lachnosterna Hope, 1837:99; type: Melolonthafer-
vida Fabricius, by original designation.
Trichesthes Erichson, 1847:658; type: Melolontha
pilosicollis Knoch, by monotypy (=tristis).
Trichestes Blanchard, 1850:141; type: Melolontha
pilosicollis Knoch tristisis). Saylor (1942:162)
cited this as an "unnecessary emendation of
Trichesthes Erichson", but Riley (1988:26)
called it a "a justified emendation."
Endrosa LeConte, 1856:234; type: Melolontha quer-
cu Knoch, by subsequent designation of Saylor
(1942).
Gynnis LeConte, 1856:262; type: Gynnis debilis
LeConte, by monotypy.
Subgenera
Phytalus Erichson, 1847:658; type: Melolontha pu-
bereus Mannerheim, designated by Saylor
(1942).
Chlaenobia Blanchard, 1850:116; type: Chlaenobia
ciliatipes Blanchard, by monotypy.
Listrochelus Blanchard, 1850:141; type: Listrochelus
laportei Blanchard, by monotypy.
Tostegoptera Blanchard, 1850:149; type: Melolontha
lanceolata Say, by monolypy.
Eugastra LeConte, 1856:233; type: Tostegoptera
cribrosa LeConte, designated by Saylor (1942).

Chirodines Bates, 1888:169;type: Chirodineszunilen-
sis Bates, by monotypy.
Triodonyx Saylor, 1942:158; type: Phyllophaga gi-
gantissima Saylor, by original designation.
Cnemarachis Saylor, 1942:159; type: Lachnosterna
andinei Smyth, by original designation. Syno-
nyms: Abcrana Saylor, 1942:159, and Clem-
ora Saylor, 1942:159-160.

Discussion: Because of the nature of the present
publication (faunal rather than monographic), it is not
appropriate to devote more space to the synonymy
than the basics listed above. More complete and
detailed accounts may be found in Saylor (1942),
Sanderson (1951),and Moron(1986). Thegenus name
Stenothorax Harris, 1827 (not 1826), has been listed by
some authors (e.g.. Moron, 1986; Riley, 1988) as a
synonym of Phyllophaga, but I have found no justifi-
cation for such treatment. The name was proposed in
the same footnote as was Phyllophaga, both being
coined for parts of the old genus Melolontha. How-
ever, Harris (1827:8) clearly stated that "Stenothorax
would be a good name for the subgenus having the
subspinosa for its type." (Riley, 1988:26 stated:
"Type species not designated"). The subspinosa,
referred to by Harris, is currently placed in the genus
Macrodactylus and has nothing to do with Phylloph-
aga.

Morphology

The dictionary definition of morphology involves
the form and structure of plants and animals. As such
it encompasses just about anything we can see on an
organism. Someone once said (and rightfully so) that
no species has been thoroughly or completely de-
scribed. The job of the taxonomist is to determine
which characters are important, of the thousands pos-
sessed by agroupoforganisms,andthenapply them to
the scheme of classification and to comparative stud-
ies.
Phyllophaga species are so similar in general
appearance that the normal comment of a casual ob-
server is: "they all look alike". That is only superfi-
cial, and the taxonomist is fortunate that excellent
distinguishing morphological features are found in
nearly all the species. The most distinctive structures
(the genitalia) happen to be hidden inside the abdo-
men, but they are extracted easily. Most of these have
been mentioned already in the Format of Presentation
and Taxonomy sections, and they are shown graphi-
cally in Table 8.

It is virtualy impossible i desembe mhe while
diffeivi,.. in innernal morphology and the Complex
genitalia. Therefore we -11 devo- only small
amount of spaw i the subject, briefy clarifying same
of die c hanxiera used.

Mean. and antatheracic fibiam with 2 spires. Taredl

The Be... Phyl~phgso is delineated by the fol-
lowing elammust"'c"~ Manadblr, easureale fro
dranad view. Leat abdominal spi-ales on appear sle.
roo-d p... oof a--.. Amen- "'now -a-.,use
[I.Mtend. Side pieev of an r-rrow Ameminal
esth 1 1 1'

one anivable in olh,,; Woth movable in females.
Sameof hlb tAheell ad in i. ky ald de.
-famans require additional cl iftestin:

ments. ber asn the bawbl scap, and mie cl ,......
Lines im, basd and no movable Howeverstrysimmee
at ..go ffirreof should be -osid-rd . dJivision
when commig segments. Pwarlosenpoons hvcyedied
heavily he length of the also in relimem 1 th de stern
(=fnlhlP), but wlahmaIpmecix paiementa. Al.
though we haw included this i the divicipifamr, we
:mwe trouble drxcidfiag whether the club is equal in
rlll to the (often l called sulbual in the lema-
pare, when almost is meant)
II A I.I pI III IA 1111A

an indernation at the mmHle (errring ..lon)-o itimut
ell od to saelsome). The clypeus may be variouly
p iniale, seIane,rug.seA A ll Ialsom; A i ,A I may be

U

.9-ants: of bhe farrole usually con-nvc, -11, no
spft l modification oftlhe ultima (Buh) or pared-
mnate (7th) sterances Malca f- hay, both of thes
iiddre mdifiidd diifdit dRh d ipesos ane iftd ..p-
ftif.. 1ares 889) ft.. diftndd
ealy -k-,er (bcf- 1889) as &l~ pronary dcan-
guishingtharacters. The st-, fir dhmuallgbdmi-

red segracres) ari-l, "oumscad by different
authors. Lugodill and Pslra (1953),-sd "S...a.
8" for the last. and Riley (1988) numbend the .,,his
squients ft.. I to 6 ( ao). Tbis lai.ology is the
.sellt of the first 2 segannia b..g hidden under it.e
ancendione, it cc al plaes. Although in= are son,
ex~len cclfcal speificttrr on the last 2 at !,,a,,
they the dli can to de-cbe, and genital ic ch arecters
_.wfrrpiabla =it tsiarT sat.
by aii-n~ichia (fig. 429431) crocang sidiffrachon g
infing Itofter app-d vlvty and sonatianci irides-
cant and can be abraded. Mh smac (o, costac are
.Wrmly reduced or barely hinficatedi, except for the
.M.] can, fig. 41641617 fr -rapriso.)
of xr" I spoc-liti-e he,,nateri-ribi-idihone spo
mj I . , "
,pper (oino), Mosth A-,ay is anovable and usually
the longer of fin 2* Th, 1-e spin rarely is codifled
drastically (e.g.,fis"Is, fig. 39S) ard a th, longerone.
Othe, Iahich -ay be confusing ase "moyabl,"
hamultd) and "fi-cd" (-unoticcnlaud). When
thespinis fixed (only in the 10,) ithas socket
ah..ing, .nd it is ino-vable. In = rig this ch--e
.bk. An.xhcr clarratener. yhich aray be nori useful

fa 422 PhyNo p . a - oad~aburb- dy- II
iiising '.(7mm-m i u )I

spor h,,h id-C ha ari b,-, aurckorand r- arphrlay ,k,, finr less ob-ii, ch--ctr Only ,
able. Arimlher sons.-er whioh may be = useful few(a.g.,therfird ... plea) formiesoannotbe dsn
..thn Uo ii -hoisly, is ihe fringe of on h lle giashed -ept by -isoiatinin iih the maler. Win.r
.rio. ii~ffio ini mulose, ammilly rforrod to as the nami.y. good to-m.logy lb. nor homi dl-elpinl
"fiffige of spioules" (fig. 425). Differen-e in the for the subrous genitalia forms, and this complicasoe
number of those Eea eetaloilated for shr-.ri, any ... bill d,,dp no The asins uisi heare inise
pmopoe by Bovmg (1942), cgln the( morml-
i maj i .- -i .~ ~ ogyand homology said atention
dismady, [ hat ,ast am be cn e =ogoeiied st at The large rale orga is composed of 3 primary
gh"nc, h gooM dhoamn... =ar aublo The elereenoi~fig.432): I)tBaslpi-F, cylarid-Ld-on
female gentadinarenearly as distinct-, ~but their gets (specallysomperloaga), curveddowtoado

posucriorend. The anterior losed byamembrane
99222 gi 29222992 ("922 9 9Dor llyi2
is called "aseram" and 'he lower par called -,ranal
ancbralnne" (behind which -eposes the invagirmacd
acdca929s). 2)Clasprs: Most a 1992ic9charactr
am found her. They ane omposed of the"phalio-
base" whichs9999ndisth Iecanhe "929 99and
the -parawases"which are theairedprolongation
often extensively modified and Cubac symm-call or
asymoeical. 3) alea9us: The inummitte organ
(=p-9 is -N 2, Wood, 1952)29. i99agim999 im9na9
The stru29 rar91yhasbeen9929-A 29nousam2ly,

shtth edengus may be much =in useful in distan-
gmshdlowely rlatedspecie We h-,illustrated
sn92ul here (fig. 433-50, S83,921,602) W show the
In- and .- 9 2s charac29rs9 th9 2n. These ware
possible only because of itd-aces in teelatiues, an
armicaltyprinadryth in 3Ams-imal fomsfea and the
u of the SEM to study itd ill.-tat the. se
121211y229y99999 d,9999992292999

K -~

difficti tidescibe. Theyd-meregmatcrautermomm
the fiam-
The female Fenitali, always are symancidca] and

mnfe-o plift, (Insial), 2) sufac- pl-te (,pp,,), and
3)apubic mess. The22plates often fused,with
a, withom. fold, or Saw= to indicate the division, and
then, shape can be meafified by dissection and dance.
risen M Eosmadron of the softer connecting am-
bmee. A, "imernal process", found on the inferior
plate at the, suddle, pmjmtsa toward the pubic proses

speies,(Youngs.fi f ig))ml9fg. 367; obso9 a, fig.
335) appear i have the pubic process abML

r..--dV j
'' VII

Key to Florida Genera
of the tribe Melolonthini (Adults)
(moldified from Howden, 1968, and Hardy, 1974)

Pyg-dum large, nrowed to a rounded .e,,

less than 12mm) ........................ D polw-as

2(1). Amernal club of 3 segments in both sexes;
elyanv2 clothed with scale, alum gltdou

men4( S elye often clothd with als, the
paten in stripes or blotch ................... 3

3(2). Ely. with scales; segments of maleantcnnal
club flattened, umred outward; male pIste-
n temi noat or barely enlarged............

3'. . .. .. ..
segments of male antennal club centered.
slightly curved outward: male posterior fe-
mur noticeably enlimged; a single endemic
Floruda species (pubescea (Cartwnghl)) ..
......................................-.. P olyl4 a

4(2). Side piece of metathorax wide: tars laws
unequal, front and middle claws with umque
Ing,, 4 al, banal lobe on the outer claw and
normal on the other: anteor tibae bldentate

4'. Sid. pieccefmetath4o4 narrw;tasal claws
similar, toothed or cleft, but never with a
basal lobe on one, alterOr tibiae moally
trdentotenm bath sies; mntum ulage,quad-
rote, largely setose: labial palpi widely wpi-
rated, placed n4 4outer apical angles: 54
4onda4species ..... Phyllophag

Biology and Behavior

We hoovcrombuied th-bjbectsheriboc .... thy
are so interrelated, and ae have devomod minimal -m
to these artas, E-ey nap-c of,, .iminals existence

mili bchioior in d-ail u, lyond da, -pe o this
pal-ri ln'. .vc ifnclfdcd onau Imcf obenlanoos,
as we]l as ciaions for mutherr aftocre.e
apeLife Hi=t: The Iii- hf, y'l,, far Florida
44 are tor44444d I4 Table 4. W4 In-4444
444 4'., spent4, time44 44 siong the4 4 s 44 44444 44
for 44& ), and flain cycl may be different under
Florida culdromi (11 tey me m mho, parts f the
camary), Tharaft-e, thedaminmablc4 should Is, uni
with i...n

each 'p"" mcnanatmia). Bmd,, arnae on falige at
night, returning to their bidig place, (mn thc -Ii add

- ' a

56

Table 4. Life cycle summary of Florida Phyllophaga.

Species State Cycle in Years' Pupation Time Citation
anxia Nebraska 3+ July Jarvis, 1966
apicata Texas 1 September Reinhard, 1941
bruneri Florida -1? winter? present work
crenulata Indiana 3,4 July Davis, 1916a
Kansas 2 August Hayes, 1925
Kentucky 3,2 summer Ritcher, 1940
Michigan 2, 3 fall Yeager, 1950
ephilida Indiana 3,2 spring Davis, 1916a
Kentucky 2 June Ritcher, 1940
forsteri ? 2,3? late summer & fall Yeager, 1950
futilis Indiana 3 July Davis, 1916a
Kansas 2 Hayes, 1920
Kentucky 2,3 summer Ritcher, 1940
Michigan 2, 3 fall Yeager, 1950
gracilis Indiana spring Davis, 1916a
Michigan 2,3 spring Yeager, 1950
hirticula Indiana 3 July Davis, 1916a
Kentucky 3,2 Ritcher, 1940
(var. comosa) Kansas 3 summer Hayes, 1920
ilicis Indiana 3 July Davis, 1916a
Michigan? fall Yeager, 1940
implicita Kansas 2,3 Hayes,1920
Indiana 3,2 July Davis, 1916a
luctuosa ? 2,3 late summer & fall Yeager, 1950
obsoleta Costa Rica, El Salvador 1 January-February King, 1984
prununculina South Carolina 2-3? Yeager, 1950
quercus Indiana spring Davis, 1916a
tristis Indiana 2 before September Davis, 1913b
Kansas 2,1 August Hayes, 1925
Kentucky 2 summer Ritcher, 1940
Michigan 2-3 fall Yeager, 1950
Texas' 1 August Reinhard, 1941
young Florida winter present work

'Most frequent time given first.
2Eggs from Hope, Arkansas.

435-

.,,, -~~-- :---

684 6-3 P yap aai lon sdau.~lrgmn f 37 ,4,3 BImO mm 16l 1 1~d~l i 33 18

43 4+0 I

g 5 'Phphg m" a m hom..,

"' "' ":"
o.smmi.

edeb cbs) jest befom da-un Fc.,I b-, nWnt
the sell a fe, inches deep to dopesit eggs singly in a
.11,. The eggs a hi. .1=rammlrd, el..g..,.

d,' befm hM~u ~ hii g .. ally i", 30l

M& 71

-. It I ... J --

days). Lmen lohve 3 olts of -bleb duration, the
thied firstarpossalsing good assamenic chimmilem (the

Lhemighthemanspeormiteeponet Thmyfeed-nlive
rams (and apparently dead organic ramonal), him,
causing damage w444d. The 4ime of pupaton is
g .. rally amifoo, for onel, spec,,,, ,wtly in the fail,
44 (4atimea (.g., 4 4erc) inesprng The entire
443le ninges f3..4I to 4 years, vk4h same 44 i
"in-itn 2 or 3 years lon, he mine egg monors.
Subtropical onditimis may speed up the4444444
444d species hi4h require several years in noml34
efinias may need only a year in 444 id44
We have hot noticed mny indication of monotte

year abundance in our dam, although these data were
not ained orroredMwith this in mind. In eorthem
msies, worker often have rfcrrcd 0 "moods" for
thos havy, alternate year emeigences (e.g., Davls,
191;Fbs,1916) Theuseofthetermo"brood"-ws

dm (1963) deftned"brood" as all the May beetes that
occrr in any one year, regardless of the number of
spe4iesinvolved. Hel sed his "bodA, BC" on the
bas, assumption that a 3-yaa, cycle was no"al, d
thus the "broods" apearcd very 3 years. This i no-
dhe normally accepted use of the form.
Adult Diel Activity: Spmies of Phyllophaga are
essentially ncitrnal, flying from dusk and aftm sun-
down to their host rees, and normally returuningt hide
ust before dawn. During o.. wudies e puzzled over

5~?"

; - ""' ~ .~.~ - -..
~ '''' ~ ' '"': '"' ~'' "'~

the taxonomic status of the sympatric, closely related
elizoria and okeechobea. After we finally sorted the
specimens to our satisfaction, we noticed that the label
data suggested a daytime activity for okeechobea
(none at light) and a nocturnal one for elizoria (nearly
allatlight). Although furtherobservations areneeded,
we believe that okeechobea is diurnal, and that this is
the species isolating mechanism that maintains the 2 in
the same locality (e.g., Archbold Biological Station).
Although there are probably some species of Phyllo-
phaga occasionally found in the daytime, we know of
only one other (Triodonyx n. sp., from Arizona; W. E.
Warner, pers. com.) which flies regularly during the
day. However, many of the relictual Melolonthinae
appear to be diurnal (e.g., Gronocarus, Hypothyce,
Phobetus, Polylamina, Thyce).
Attraction: May beetles obviously are attracted
to their host plants and to their mates. The nature of
these attractions (possible sex or aggregating phero-
mones) has received little attention, and we have not
studied them. However, the most notable attraction is
that to light (especially ultraviolet). This behavior has
produced nearly all of our specimens for study. Their
attraction to light was known to early workers as well,
and led to the control recommendation of suspending
a light over a tub of water with a layer of kerosene on
top. The nature of the attraction is still poorly under-
stood, although types of lights and height of light trap
location were studied by Chandler, et al. (1956). The
nature of the light attraction was the subject of a 10-
paper symposium (USDA, 1961), and various trap
designs were discussed by Hollingsworth,etal. (1963).
I have found that even the flightless cribrosa group are
attracted to lights in Texas.
Season: In northern states, most species appear in
May or June, giving rise to the common name. Even
though this generality holds for most species in many
areas, some species emerge in the fall, especially in the
desert areas of the southwestern United States. The
season often appears to be extended (both early and
late) in Florida, as can be seen in the records undereach
species. We have collected brunerievery monthofthe
year in Miami, but it appears to have 2 population
peaks (one in May-June, one in Aug-Sept.) and possi-
bly 2 generations per year.
Plant Hosts: Adult beetles feed on a great variety
of plant species in diverse plant families (see species
treatments under the heading "Adult Host Plants").
Most are found on broad leaved trees and shrubs, buta
few are primarily conifer feeders (e.g.,prununculina,
debilis, tacodii, and clypeata). The latter 2 species
appear to be host specific to cypress. Of those feeding

on the non conifers, they have such diverse hosts
(sometimes with different preferences in different
localities) that few generalities can be made. I studied
the negative and positive hosts for the introduced
bruneri in Miami (Woodruff, 1961) without finding
any pattern of plant families. Sometimes a close
relative (in the same genus) of good host would have
no beetles or feeding signs. The positive hosts and
their families are shown under bruneri in the species
treatments.
General Behavior: Other aspects not heated above,
but which had some discussion in the literature, in-
clude: proportion of the sexes (Chamberlin & Seaton,
1941); larval spatial pattern (Guppy & Harcourt, 1970);
oviposition (Sweetman, 1927); height of flight (Sh-
enefelt & Simkover, 1951); mite symbiosis (Jarvis,
1964); soil temperature influence on grubs (McCol-
loch & Hayes, 1923; species of a single host (haw-
thorn) (McCulloch & Hayes, 1922); and especially in
the following general references: Chamberlin, et al.,
1943-1941; Forbes, 1891b-1916; Hayes, 1925, 1929;
Hayes & McColloch, 1928; Ritcher, 1939-1958.
Immature Stages: Unfortunately we know less
than half the larvae for our Florida species, and only
the pupa of bruneri has been described (Woodruff,
1961). The known larvae are summarized in Table 4,
and they are treated in the following key. Taxonomic
characters are found primarily on the last ventral
abdominal segment (fig. 460) and the epipharynx (fig.
459). Space does notpermit a moredetailed treatment
here, but theknown larvae are described in detail under
each species. The primary references to larvae are as
follows: Boving (1937, 1942); Boving & Craighead
(1931); Davis (1913-1920); Hayes (1925-1929); Hayes
& McColloch (1920, 1928); Moron (1958); Peterson
(1951); Ritcher (1938-1967); Uhler (1941); Wade
(1935).
Phyllophaga pupae have rarely been treated, and
never in a taxonomic sense. Ritcher (1939) reviewed
the literature on the depth at which pupation takes
place. He found that some species (e.g., hirticula)
pupated deep in the soil, others (e.g., inversa) were
shallow, and most were intermediate (e.g., futilis).
Factors influencingdepth included the structure, drain-
age, and moisture of the soil,but they "did not change
the relative positions of the various species." Differ-
ences in latitude also had no effect on the depth for a
given species.
Moron (1986:163-168, fig. 281-287) described
and illustrated a typical pupa, the spiracles, and sexual
characters for a Mexican species (brevidens Bates).

fig. 457. Phyllophaga brunertr third instar larva (13mm 1.5mm).

fig. 459. Phyllophaga bruneri: epipharynx of third instar larva.

,Anal Slit

fig. 458. Phyllophaga bruneri: head of third instar larva, dorsal.

fig. 460. Phyllophaga sp.: ventral view of 10th abdominal seg-
ment with raster, diagrammatic (After Boving, 1942). Brb =
barbula; LAL lower anal lip; Pa palidium; Prsept preseptular
setae; Sept septula; Tgl = tegillum; UVA upper anal lip.

KEY TO KNOWN LARVAE 6(2). Proplegmata absent; pali 10-13; dorso-exte-
OFFLORIDA PHYLLOPHArGA triorregion of mandible with no punctures; pali
OF FLORDA PHYLLOPHAGA depressed, short, bent toward septula; septula
(modified from Boving 1942 and Ritcher 1966) broadly oval; crepidal punctures 6 or less
(probably includes apicata also) ................
.................................... tristis (Fabricius
1. Palidiaabsent(Fig. 477)(subgenus Cnemra- 6. Proplegmata present, at least on 1 side of
chis) .................................. bruneri Chapin epipharynx ........................................ 7
1'. Palidiapresent(Fig.460)(subgeneraPhytalus
& Phyllophaga) .................................... 2 7(6') Proplegmata either indistinct and often differ-
ent in number on both sides, or not more than
2(1). Maxillary articulating skin ventrally with short, 4; dorso-exterior punctures of mandible (nor-
conical, orrounded dark granules; pali usually mally) present; pali hooked, 12 or more, or
straight, pointed tips (hooked in quercus) .. long sharply pointed and 20 or more ........8
......................................................... 3 7'. Proplegmata distinct, usually same number on
2'. Maxillary articulating skin ventrally with long both sides, more than 4; dorso-exterior punc-
and short setae, and sometimes with small, tures of mandible absent ................ 12
ring-shaped pale punctures among the setae,
but never with thick, dark granules; pali vari- 8(7). Proplegmata 25 or more ...................
ous shapes................................. ............. 6 ........................ prununculina (Burmeister)
8'. Proplegmata 20 or less ..........9............... 9
3(2). Proplegmata distinct, 7 to 14, or a few more;
pali sometimes curved, with straight lateral 9(8). Proplegmata about 20 .... debilis (LeConte)
edges and sharp points; numbering 8-27; pre- 9'. Proplegmata 10 or less .... ................. 10
septular setae numerous; and lobes with many
pointed short and strong setae .................. 4 10(9). Pali 17-20; proplegmata 6 or 7 ..................
3'. Proplegmata absent or vestigial, appearing ............................ glaberrima (Blanchard)
faintly as several fine lines on each side; pali 10. Pali 20-25 or more; proplegmata 8-10 .... 11
10-20 or a few more; maxillary articulating
skin ventrally with 9-35 or more granules .. 11(10'). Dorso-molar region with about 12 setae and
........................................ quercus (Knoch) fewer than 10 punctures in front of them at
inner margin of scissorial part ....................
4(3). Maxillary articulating skin with short, thick, ................................. ephilida (Say)
pointed granules; palidium with 1 irregular 11. Dorso-molar region with 10-15 punctures ..
row of 25-27 pali; proplegmata 7-10 .......... ........................................ utilis (LeConte)
........................... ............ aemula (Horn)
4'. Maxillary articulating skin with many more 12(75. Pali 9-10; dorso-exterior region of mandible
than 5 granules; palidium with 18-24 pali; pro- with no punctures .......................... 16
plegmata 12-15 ........................................ 5 12'. Pali 12-30; dorso-exterior region of mandible
5(4. Maxillary articulating skin with 14-21, some- with 0-30 punctures ................................ 13
5(49. Maxillary articulating skin with 14-21, some-
times more, cone-shaped granules; heli 8-10; 13(12). Pali 20-30 ............. ... .............. 14
proplegmata 12-15; palidium with 20-24 or 13'. Pali 12-18 .................................... 16
more straight pali ...... crenulata (Froelich)
5'. Maxillary articulating skin with 25-35 cone- 14(13). Proplegmata 11-15; preseptular setae 6 or
shaped granules; heli 14; proplegmata ap- more; pali 20-30; dorso-exterior region of
proximately 15; palidium with about 18 some- mandible with either 3-10 or 20-25 punctures;
what curved pali .............................. .. pali in irregular row (some parts double)....
.................................. parvidens (LeConte) ....................................... anxia (LeConte)

14'. Proplegmata 6 or less; preseptular setae absent
(latifrons) or 7-10 (micans); pali 22 or more;
dorso-exterior region of mandible with either
few (1, 2, or absent) to 15 (micans) or 3-10
(latifrons)........................... ............. 15

15(14'). Proplegmata4 or less; pali 22or a few more in
1 quite regular row and closely set in whole
palidium; dorso-exterior region of mandible
with few (1-2 or absent) to about 15; scrobis
with 8 or fewer punctures; preseptular setae
absent .(probably includes cupuliformis also)
........................................ micans (Knoch)
15'. Proplegmata 4-6; pali about 26, palidium ex-
tending forward with several (usually 3-5) pali
in front of tegillum; dorso-exterior region of
mandible with 3-10 punctures; scrobis with 8-
18 punctures; often 1-2 preseptular setae ..
............................a... latifrons (LeConte)

16(13'). Dorso-exterior punctures ofmandible 1-4;pali
13-16; proplegmata 3 or less, short, weak, not
always same on each side; preseptular setae
absent ........................ dispar (Burmeister)
16'. Dorso-exterior punctures of mandible 15-30;
pali 12-18; proplegmata 5-8 on one side, less
than 5 on other; preseptular setae normally
absent, but many specimens with 1-5 ........
...................... ........ gracilis (Burmeister)

17(12). Both right and left chaetoparia with numerous
punctures among the setae, even spreading
into pedium .................... luctuosa (Horn)
17'. Only right chaetoparia with numerous punc-
tures among the setae............................ 18

18(17'). Dorso-exterior region of mandible with 10 or
less punctures ................................... ... 19
18'. Dorso-exterior region of mandible with 18 or
more punctures .................................... 20

19(18). Pali about twice as long as wide at base, vary-
ing in size in different parts of palidium, very
.closely set and irregularly arranged; dorso-
exterior region of mandible with 5 or less
punctures ............................ ilicis (Knoch)
19'. Pali at least 3 times as long as wide at base,
about same size throughout palidium, less
close than in ilicis and in an almost regular
row; dorso-exterior region of mandible with
10 or fewer punctures ............................
............................ profunda (Blanchard)

20(18). Dorso-exterior region of mandible with 20 or
less punctures ................................. ... 21
20'. Dorso-exterior region of mandible with 30 or
more punctures.................................. 22

21(20). Proplegmata generally long and curved, usu-
ally 9-11; septula elongate, subelliptical, slightly
constricted in middle; preseptular setae 4 or
less; pali 25-35 .................... hornii (Smith)
21'. Proplegmata fairly short and almost straight,
usually 7-8; septula subrectangular; preseptu-
lar setae about 7; pali 27-32 ..................
.................................. forsteri (Burmeister)

22(20'). Pali 20; proplegmata 10; preseptular setae 5;
crepidal punctures about 50; scrobis with a
longitudial series of about 10 punctures and
about 6 setae .... knochii (Schoen. & Gyll.)
22'. Pali about 30; proplegmata 8-10; preseptular
setae 6-8; crepidal punctures about 35; scrobis
with a longitudinal row of about 6 punctures
and 1 to 5 setae................ hirticula (Knoch)

fig. 461. Phyllophaga bruneri: pupa, ventral (25mm 3mm).

Distribution and Zoogeography

Although different names have been used for
Phyllophaga in various parts of the World, we feel that
nothing is accomplished (and much is lost) by splitting
the group into genera purely on the basis of geography.
We therefore believe that Phyllophaga (sensu lato)
occurs in both Eastern and Western Hemispheres,
even though the European and Asiatic literature often
uses the names Lachnosterna, Holotrichia, and Brah-
mina.
With the above as a premise, we believe there are
probably more than 1000 species of Phyllophaga. We
have not studied Old World species, and little further

will be said about them here. In the New World, about
200 species are found in the United States, 254 in
Mexico (Moron, 1986), 80 in South American (Frey,
1975), and probably more than 150 in the West Indies
(Sanderson, pers. com. and Woodruff, unpublished).
Several checklists for the United States were
consulted for various reasons, but their short (and often
vague) distribution records were of limited use. However,
they are listed here for the benefit of the reader: Austin
(1880); Blackwelder (1939, 1957); Blackwelder &
Blackwelder (1948); Crotch (1874); Dalla Tone (1912);
Haldeman & LeConte (1853); Henshaw (1885, 1887,
1889, 1895); LeConte (1863); Leng (1920); Leng &
Mutchler (1927, 1933).

Table 5. Alphabetical list and references to known larvae of Florida Phyllophaga.

Hayes, 1925 Hayes, 1929

Boving, 1942

Ritcher, 1966

p.7

p. 6

[

p. 5

aemula
anxia
apicata*
bruneri
crenulata
cupuliformis*
debills
dispar
ephilida
forsteri
futills
glaberrima
gracills
hirticula
hornll
ilicis
implicita
knochii
latifrons
luctuosa
micans*
obsoleta
parvidens
profunda
prununculina
quercus
tristis*

p. 29-30; fig. 2-8
p. 45; fig. 114-116 p. 86-87
0-72 fig. 27,188 p. 33; fig. 33-36 p. 86, 87, 97; fig. 225
[Woodruff, 1961: 1-31; fig. 1-26]
1-63 fig. 185 p. 30; fig. 9-13 p. 86,88,91,93; fig. 18
fig. 170 p. 42; fig. 106-109
p. 36; fig. 63-65
p. 40-41; fig. 92-94
fig. 161 p. 40-41; fig. 83-85 p. 86-87, 89; fig. 220
Hayes, 1920: 307] p. 53; fig. 199-203
fig. 37, 69,164 p. 38; fig. 72-73 p. 86-87, 89; fig. 219
p. 39; fig. 86-87
fig. 163 p. 40-41; fig. 99-101
6-58 fig. 182 p. 51; fig. 179-182 p. 86-87, 91, 95, 97; fi
fig. 173 p. 52; fig. 191-194 p. 86, 88, 91,93,95; fi
fig. 184 p. 50; fig. 168-172
fig. 180 p. 45-46; fig. 133-137 p. 86-87
p. 51; fig. 183-186
fig. 160 p. 40; fig. 88-91
p. 47; fig. 148-151
fig. 170 p. 42; fig. 106-109 p. 86-87, 93; fig. 201
[King, 1984: key p. 45-48; fig. 7]
p. 31; fig. 14-17
fig. 179 p. 50-51; fig. 173-176 p. 88, 95; fig. 208
p. 38-39; fig. 79-81
p. 31; fig. 18-22 p. 87
0-72 fig. 27,188 p. 33; fig. 33-36 p. 86, 87, 97; fig. 225

0, 198

g.184,205,207,217,224
ig. 176, 188, 189, 204, 213

*P. cupuliformis/micans and apicata/tristis adults are so similar that larvae are indistinguishable at present.

p. 7

Within the United States, there have been many
state lists or faunal studies which give a general idea
of Phyllophaga species found therein. These are
variable in quality of both the collecting and the
knowledge at the time they were prepared. They are
listed alphabetically by state, with the number of
species in brackets, and with the author and date in
parentheses: Alabama [66] (Loding, 1945); Arkan-
sas [44] (Sanderson, 1944); Arizona [44] (Butler &
Wemer, 1961); Connecticut [13] (Britton, 1920);
Florida [42] (Young & Thames, 1949), [54] (present
study); Georgia [66] (Fattig, 1944); Illinois [34]
(Forbes, 1916); Indiana [26] (Blatchley, 1910); Iowa
[33] (Travis, 1934); Kansas [44] (Knaus, 1897);
Kentucky [36] (Ritcher, 1940); Louisiana [62] (Riley,
1988); Missisppi [45] (Langston, 1927b), [53] (Lago,
1980); Missouri [27] (Owens, 1950); Nebraska [25]
(Dawson, 1922), [+8] (Ratcliffe, 1974); New Jersey
[32] (Smith, 1910); New York [30] (Leonard, 1926);
North Carolina [39] (Brimley, 1938); North Dakota
[10] (Lago, et al. 1979); Ohio [36] (Neiswander,
1963); South Carolina [31] (Luginbill, 1928) [55]
(Kirk, 1969, 1970); South Dakota [17] (Kirk &
Balsbaugh, 1975); Texas [96] (Reinhard, 1950); Wis-
consin [18] (Chamberlin, et al., 1943)
We have attempted to provide detailed distribu-
tion records for Florida and also the United States
distribution for those recorded here. These are dis-
cussed under the DISTRIBUTION heading, and maps
are provided for Florida (the dots as near the localities
as possible) and the U. S. (general distribution area
outlined and indicated by shading; questionable rec-
ords are so indicated). We used Luginbill and Painter
(1953) as our basic source, but their maps and written
records were not documented by citations. Theirdots
usually appeared in the center of a state, although this
is misleading in several coastal species in the south-
east. We have tried to verify any unusual distribution
records, butwehavenot alwayssucceeded. Although
most of the major state lists are noted in the previous
paragraph, there are many smaller references, dealing
with only a few species. The several papers by Hatch
(1928,1929, 1941) are extremely useful as a starting
point for such a literature search.
In the Florida fauna, 3 species (bruneri, pube-
rula, andyoungi) are introduced from the WestIndies.
P. obsoleta has basically a Mexico-Central America-
South America distribution, just barely extending into
the United States, in Florida occurring only in the
panhandle. The following 8 species are recorded only

from Florida and may be precinctive (endemic): elizoria
Saylor, elongara Linell,floridana Robinson, okeechobea
Robinson, ovalis Cartwright, skelleyi n. sp., and tecta
Cartwright
Other New World references, by geographic area,
which may be useful in interpreting zoogeography of
Phyllophaga are: Central America (King, 1984; King
& Saunders, 1979, 1984); Cuba (Garcia-Vidal, 1975-
1987); Hispaniola (Sanderson, 1951); Lesser Antilles
(Chalumeau, 1983,1985;Chalumeau&Gruner, 1976);
Mexico (Moron, 1986); Puerto Rico (Wolcott, 1948);
South America (Frey, 1975); andWestIndies (Saylor,
1940).

Economic Importance

Few other groups of insects are as abundant and as
economic in both the larval and adult stages as are the
"white grubs" and "June bugs". Although we have
seen severe damage to sod and tree foliage in Florida,
ithasnotbeen asextensive asreportedelsewhere. Most
references discuss economic damage, but few have
quantified it. In addition to isolated economic reports
and those of a general nature, we have tried to summa-
rize, in the following paragraphs, some references with
meaningful figures.
Surprisingly,Luginbilland Painter(1953)devoted
little space to economics in their treatment of the U. S.
species. They mentioned that thousands of acres of
pasture in a single locality had the feed value reduced
by larvae, and the damage of white grubs to corn,
timothy, and potatoes (during peak years in the north-
central states) amounted to "... several million dollars
in a single year." They suggested that one reason, that
damage by adults in the south is seldom serious, is the
"... steady rise in temperature in the spring ... causes the
adult May beetles to emerge gradually."
Davis (1916a:270) gave more graphic data on
numbers. In May and June of 1914, he surveyed the
area of northern Illinois, Iowa, Wisconsin, and Michi-
gan, where he found "... the timber, which consisted
chiefly of oak and hickory, was completely stripped of
its foliage ... the beetles were so abundant that the dead
ones accumulating beneath the lights had to be swept
away each morning to prevent or at least modify the
terrible stench which they produced." The volume was
so great that "At one small town in Wisconsin the
beetles accumulating beneath the ten arc lights of the
town were hauled away each morning for period often
days or two weeks, by the wagon load."

In 1918, Davis provided more specific damage
figures for the heavy larval infestations of 1912 in
Iowa, Illinois, and Wisconsin. He estimated the damage
to corn, timothy, and potatoes in that area at $7,000,000,
and in other areas at $5,000,000. He also gave esti-
mates of larval populations in heavily infested areas at
106,680 per acre. In the fall (when they are well-fed),
he estimated that each grub weighed 1 gram (454 per
pound), and thus there were 235 pounds of grubs per
acre. He even recommended feeding them to hogs
because they had nearly the samepercentage offat and
protein per pound as corn (although deficient in carbo-
hydrates). Hisfigureoffoodvaluebeinggood at$3.00
per acre (presumably 235 pounds) is related to the
economics of the time, and can hardly be compared
with modem costs.
Forbes (1908:166) had earlier given statistics on
the quantities of grubs found in a 10-acre destroyed
cornfield in Illinois. By sifting the soil of each hill (3.5
ft. square) to a depth of 20 inches, he collected 3460
grubs per 100 hills, or "... about three hundred pounds
ofgrubstotheacre." Swine (100pigsabout75pounds
each and 8 large sows) were introduced into the field
on September 23, and by October 13 (20 days) the
entire 10 acres were rooted to a depth of 10 to 12
inches, and the grubs were reduced by 86% to4.8 per
hill. He did point out that the thorny-headed worm of
swine (Echinorhynchus gigas) uses white grubs as an
intermediate host, so that swine should not be used for
control where they have been pastured previously.
Forbes also mentioned hand collecting of adults as a
means of control. He quoted statistics from Sweden
(presumably for the related genus Melolontha) in a
ten-year period (1885-1895), when residents collected
29,736 bushelsofbeetlesata laborcost of$15554 (52
cents per bushel). They believed that this system
reduced the population from 14,801 (1887) to 5,611
bushels (1895), even though more collectors were
involved in 1895.
Since conifers are one of the most significant
elements of Florida forests, the species feeding on
them have an economic impact, however difficult to
assess. Grubs are known to damage pine nursery stock
(Baker, 1972,1985;Nairn &Wong, 1965; Shenefelt &
Simkover, 1951; Yeager, 1950), and adults of several
species damaged pine flowers in Louisiana (McLem-
ore, 1953).
Rolston and Barlow (1980) reported larvae dam-
aging sweet potato tubers in Louisiana sufficiently to
require preplanting insecticide treatment.
In Mexico, Moron (1986:7-8) reported a reduc-
tion of 1,314 kilograms per hectare of corn, due to

Phyllophaga larvae. He listed another case in the state
of Jalisco where grubs destroyed 13,784 hectares of
corn, beans, rice, and sorghum. Damage resulted in
the use of many insecticides over the years: Carbofu-
ran, Chlorfenvinfos, Chlorpirifos, DDT, Diazinon,
Foxim, heptachlor, Isofenfos, Protiofos, and Terbufos.
The use of such a chemical arsenal has had not only an
economic impact, but a profound effect on the envi-
ronment as well.
In Florida, sugarcane is the one specific economic
field crop which may be affected by white grubs.
Although grubs of other scarab genera are pests, appar-
ently the only species of Phyllophaga involved is
latifrons (Gordon & Anderson, 1981; Cherry, 1984-
1985; Sosa, 1984). The introduced species, bruneri, is
a known sugarcane pest in Cuba(Stahl & Scaramuzza,
1929), but it has not yet reached the Lake Okeechobee
sugarcaneregion. Because many speciesofPhylloph-
aga are sugarcane pests elsewhere, the following addi-
tional citations may be useful: Box (1953); VanDine
(1926); VanDine &Christenson (1932); Wade (1951);
Williams, et al. (1969); Wilson (1969).
Wolcott(1914-1948) devoted considerable atten-
tion to the white grubs as pests of sugarcane in Puerto
Rico, where they were once an economic threat. He
stated (1948:250) that they "... reached such a peak of
abundance as to become the decisive factor in deter-
mining whether the growth of sugarcane or any other
crop was possible." He further indicated that they "...
were the major insect pest of every crop grown ... in
bulk,... overbalanced all the other insects of the Island,
not only individually, but all other insects combined."
All this was before the introduction of the giant Suri-
nam toad (Bufo marines L.) which brings us to the next
section on natural enemies.

Natural Enemies

As pointed out in the previous section, these in-
sects can occur in tremendous numbers (wagon loads!).
When such populations explode, they are often fol-
lowed by some measure of control by natural enemies,
such as predators and parasites. We all know the story
of seagulls and grasshoppers in the history of the
Mormons in Utah.
From the economic viewpoint, we are fortunate
that there is an entire complex of such natural enemies
of the genus Phyllophaga. J. J. Davis (1919), the
author of the "wagon loads" quote, published an
extensive monograph on these natural enemies. We
have mentioned most of these under the species of
beetle parasitized or preyed upon. Because chemicals

have not provided ,arisf- toy ... ai aad fuccau
ca ..... W p4, bl4ms, amid captas is being
plaed in biolgalccat 1 Space does runpassitai
f th-bjectatm lial~lesu j bid-po wprvidl
Ih. fall-g -i~esa ad falce

94194441949
Hy.414 p-. W9Ial44 s, 9 he48) ll. ag- m ears-
sirea ze be. samaed (eef-siolls in parendieses):
DIPTERA.

suid-i~r Crypwo c ge~i, hsuai, Walka,,
Cryvi. a if ... vnar Widtmn, Euro-d,,j,,,-
Is, Zinon, Eun-im mile CoqulluL Biompsa lachnos-

Rico, Wolcon, 1948).
S,,, d7. I ... 4441phlge pe9hibia Ald449h,
--d-o mrcmolr Aldnch, S. conbicur
Townsend, S. hellcis Townsend, S. unlis Aldneb, S.
944c,191, Pan.,, S.4 sp, (Davis, 1919); S. basslis
waF,, S 4bdla. ,yk., f41,S i4,,g 4141114ig,
1944).
Frou he lia'se, the foll-bign i-,F panad.te
li_ been ancdd

HYMENOPTERA:
hplol --tvi iorot Say, T_ punu-i

Daft Tom E 1 41p Sly, E 4 14
F1,,ei., (Davis. 1919): Tiphla cosps All.4, T
4n- Say, T. -194'9419 Say, T. 91gni, Rub.;
my- -Ioa-limn P-'e"r m. usb-~ C-.'n M
cula9 Fabric, M. ,obs- P9abricius91 (P9g., 1944
Tiphus alas-l Say (Riley, 1874, Petch &L Hanouni,
1P92;hul,,, 1907; WSIcoI 1914); T9ph,, bob..do
Allen (Berbat & He4449. 19704 Rivas, Mayo, .
Helsl, 1979; 9,41, 1966).
Scelod.c Capes-,iipl..dpas oayIC q.,d-

1919(P,,,g, 194L944I,&91 .499.99
(Calipo,-ui) o-if-i-ot Fabn,,.,, D d...s

1949).
P1lec99)4191 44'4'1,, 49ol44). (Davs,
1919; Fattg. 144; Lim, Yule,&g Slowart. 1980; Pachh
& H.-and, 1925).
Iclinciaronisise: Oph-io bfmovltoi, Boill,
(Forbes, 19G7; Davis, 1919); 0 aacyl- Conalicas,
0 p-,.,-( Say (Fautg, 1944).

DIPTERA:
Tachinidae: Microphthalma disjuncta Wiedeman,
M. pruinosa Coquillett (Davis, 1919); M. disjuncta
Wied., M. divisa Wied. (Fattig, 1944); M. disjuncta
Wied. (Forbes, 1907). M. phyllophagaeCurran (Petch
& Hammond, 1925).
Dexiidae: Ptilodexia harpasa Walker tibialisis
Desvoidy), P. abdominalis Desvoidy, Myocera crem-
ides Walker?, Prosena (Mochlosoma) lacertosa ven
der Wulp (Davis, 1919); Ptilodexia cerata Walker
(Fattig, 1944).
Bombyliidae: Sparnopolius fulvus Wiedeman
(Forbes, 1907; Davis, 1919; Fattig, 1944). Hyperparasite
Exoprosopa fascipennis (Forbes, 1907); E. fasciata
(Ritcher & Fluke, 1935).
Predators: The number of vertebrate animals.
known to feed on Phyllophaga, is too extensive to list
here, but involves many birds, mammals, and toads.
Although the number of insect predators is probably
much larger than recorded, the following species are
known as predators on larvae:
DIPTERA:
Asilidae: (larvae of the robber flies feed exter-
nally on white grubs): Promachus vertebratus Say, P.
Jitchii Osten-Sacken, P. bastardii Macquart, Erax macu-
latus Macquart interruptsts Macq., E. lateralis Macq.),
E. aestuans Linnaeus, E. cinerascens Bellardi (=al-
bibarbis Macq.), Deromyia winthemi Wiedeman, D.
discolor Loew, D. umbrina Loew, Asilus paropus
Walker,A. lecythus Walker, Ceraturgus cruciatusSay
(Davis, 1919); Asilus notatus Wied., A. virginicus
Banks,DeromyiadiscolorLoew,D. termataLoew,D.
winthemi Wied., Erax interruptus Linn., E. rufibarbis
Macquart, Proclacanthus brevipennis Wiedeman, P.
longus Wiedeman, P. rufus Willison, Promachus bas-
tardii Macq., P. rufipes Fabricius (Fattig, 1944).
Tabanidae (larvae of horse flies feed externally on
white grubs): Tabanus sulcifrons Macquart, T. atratus
Fabricius (Davis, 1919); T. americanus Forster, T.
fulvulus Wiedeman, T. molestus Say, T. nigrescens
P.deBeauvois (Fattig, 1944).
COLEOPTERA:
Carabidae (larval and adult ground beetles prey
on white grubs): Harpalus pennsylvanicus Dejean, H.
caliginosus Fabricius, Calosoma calidum Fabricius
(Davis, 1919); C. sayi Dejean, C. scrutator Fabricius,
Dicaelus dilatatus Say, D. purpuratus Bon., Harpalus
caliginosus Fab., H. erythropus Dejean, H. pennsyl-
vanicus Dej., Pasimachus punctatus Haldeman, Scaphi-
notus unicolor Fabricius (Fattig, 1944).
Elateridae: Pyrophorus lumiiosus Illiger (Davis,
1919; Wolcott, 1948); P. havaniensis Castelnau (pres-

ent work).
Miscellaneous. The following groups of parasites
or predators are either minor ornon-insect and are only
briefly and generally cited here.
Mites: Davis (1919:102) recorded Rhizoglyphus
phylloxerae Riley, Tyroglyphus armipes Banks, and
Parasitus sp. on larvae, and Uropoda sp. on adults.
Jarvis (1964) discussed the association between a spe-
cies of Caloglyphus (Acaridae) and Phyllophaga anxia.
It was described as a new species by Oseto and Mayo
(1975). Fetch and Hammond (1925) listed hypopial
nymphs of Tyroglyphus sp. infesting 100% of white
grubs in one area in Quebec, but they found Rhizoglyphus
phylloxerae infrequently.
Parasiic worms: The helminth known as the thorny-
headed worm of swine (Echinorhynchus gigas) was
reported by Stiles (1892) to be secondary in white
grubs. It was discussed in detail by Glasgow (1926).
Hall (1929) discussed helminths and their intermedi-
ary hosts. Davis (1919) reported only a few larvae
infested with hairworms (Merminthidae) of the genus
Mermis (one of which was obtained at Pensacola,
Florida). Davis (1919) reported 2 species of nema-
todes in white grubs: Diplogaster aerivora Cobb and
Cephalobus(?) sp. Berberet and Helms (1969) studied
2 species of Eugregarina (Gregarina sp. and Actino-
cephalus sp.) in Phyllophaga anxia.
Spiders: Davis (1919) listed the following spiders
preying on adult Phyllophaga: Lycosa helluo Wakeman,
Xysticus gulosus Keys, and Plectana stellar Hentz.
Jennings (1974) discussed crab spiders preying on
scarab beetles. In our study, we found Phyllophaga
young being consumed by Nephila clavipes (Lin-
naeus), and P. skelleyi was trapped by Acanthepeira
marion Levi.
Toads: We cannot treat natural enemies without
considering the giant Surinam toad (Bufo marinus L.)
which has been introduced around the world to control
white grubs in sugarcane. Phyllophaga guanicana
Smyth, from Puerto Rico, is possibly extinct from this
toad's predation. Wolcott (1948) treated the success in
control of white grubs in Puerto Rico. Although the
toad was introduced into south Florida many years
ago, it does not seem to be extremely abundant and has
had no apparent effect on the Phyllophaga fauna here.
Protozoa: Little information is available on the
protozoan parasites which seem to be common and
play a commensal role in Phyllophaga white grubs.
Travis and Becker (1931) described and illustrated the
following from white grubs: Class Mastigophora
(Polymastix phyllophagae n. sp., Monocercomonas
melolonthae Grassi, Eutrichomastix passale Tanabe,

Eutrichomastix phyllophagae n. sp., Embadomonas
phyllophagae n. sp.). Class Rhizopoda (Arcella sp.,
Allantion sp.).
Diseases: Many types of diseases have been found
in scarab larvae, and some are available in commercial
formulations to use in control. Milky disease is one of
theimportantones(Dutky, 1941). Micrococcusnigro-
faciens Northrup, a bacterial disease, was considered
of minor importance in control (Davis, 1919). Several
fungus diseases of white grubs have been reported.
Metarrhizum anisopliae Metsch. (the green muscar-
dinefungus).Isaria densa Link (=Botrytis tenella) (the
white muscardine fungus), and Cordyceps melolonthae
Tulasne all were reported by Davis (1919). Isaria
vexans Pettit and Cordyceps melolonthae (Tulasne)?
were recorded by Pettit (1895). In Nova Scotia, Piers
(1889) reported a fungus from May beetle larvae
(probably Cordyceps melolonthae).
General references dealing in part with parasites
and predators: Davis, 1918,1919; Fattig,1944;Forbes,
1907, 1908; Lin, 1979; Lin, Stewart, & Yule, 1981;
Lin, Yule, & Stewart, 1980; Miner, 1952; Petch &
Hammond, 1925, 1926; Wolcott, 1914,1937,1948.

QUESTIONABLE AND POTENTIAL
RECORDS

In any study of this nature, dealing with thousands
of museum specimens and voluminous literature rec-
ords, errors are likely to be encountered. We have
included here those records for which we have some
doubts, specimens which have been intercepted at
ports and are potential introductions, and those which
we believe are most likely to be added to the Florida
faunal list. These are treated below, alphabetically
under each species. In addition, we must point out
what we believe to be a number of mislabelled speci-
mens (perhaps in othergroups of insects as well), all of
which bear the collector label F. W. Walker, mostly
1920 to 1922, and from Gainesville, Florida (mostly in
the University of Michigan Museum of Zoology). A
similar case, involving the same basic label data, was
encountered for another scarab of western origin
(Canthoan imitator Brown), which resulted in the erro-
neous description of a new subspecies floridanuss
Brown) based on this disjunct distribution (Woodruff,
1973:38-39). During the present study, we saw several
specimens with this data, some of which were well out
of the normal geographic range, including the only
Florida records for ilicis and ephilida, as well as the
only Gainesville records for georgiana and hirticula.

We are convinced that the above Walker speci-
mens, and potentially any with similar data, were
mislabelled. According to Dr. T. H. Hubbell (pers.
comm.), F. W. Walker was normally meticulous about
keeping field notes, for which a number was usually
attached to the pinned specimen. In reference to the
Canthon mentioned above, Hubbell stated "I think
your surmize that they were mislabelled is probably
correct, ... the chances are that Walker had nothing to
do with this material."
In general, we have had other specimens for which
we had good evidence or good reason to doubt their
validity. We have eliminated most of these from our
data bank without comment. Our basic philosophy is
that it is much more difficult to eliminate an error in
print than it is to addor clarify questionable records by
future collection or study. Some distribution records
perpetuated by Luginbill and Painter (1953) are im-
possible to refute or check because they gave no data
source for their maps or statements.
Two other incidents point out the possibility of
disassociating genitalia from the specimens, causing
confusion and, in one case, synonymy. Luginbill and
Painter (1953) used special "smoking" technique on
the genitalia to eliminate glare during photography, so
these specimens are easily recognized. Unfortunately
they did not reassociate the genitalia with the speci-
mens. Their figures and genitalia (which we have seen
in the USNM) for floridana are actually the new
species, pseudofloridana, but we could not locate the
beetles from which the genitalia were dissected.
The other incident was even more confusing.
Sanderson (1950) described murrea from a unique
female from Florida. He mentioned that it resembled
elongatla, but that the female genitalia were of a very
different type, comparing it to congrua. No further
specimens have ever been found that match this com-
bination. We borrowed the holotype from the UMMZ
and mounted it for the scanning electron microscope
photographs. Although we treated it as a species
during preparation of this manuscript, we are now
convinced that the body is that of elongata and the
genitalia are those ofclypeata, creating the new synon-
ymy.

Individual Species Discussions

Phyllophaga analis (Burmeister)
(=bifoveolata DuVal, subsericans DuVal)

Two specimens in the FSCA represent intercep-
tions of this Cuban species at Miami, Florida: (1) 19-

III-47,R. Baker, onSolanum melongenaLinnaeus var.
esculentum Nees, from Cuba, SPB Ace. #95342; (1) 1-
VI-56, F. A. Buchanan, from Cuba.
Chapin (1932:197-199, fig. 31) redescribed and
illustrated this species, recording itfrom the following
Cuban Provinces: Havana, Santa Clara, Camaguey,
andtheIsleofPines. Healsomentionedthat a pairwas
taken at Ancon, Panama, May 10, 1909, feeding on
marineses, the only Cuban species known to him
with a continentalrecord. Box (1953)andStahl (1929)
both listed it as a pest of sugarcane. There is no
evidence that it is established in Florida.

Phyllophaga cribrosa (LeConte)

Young and Thames (1949:126) recorded a speci-
men of this species from "?Orange County, H. T.
Townsend". Although they indicated that it was "...
very doubtful", they state "It may, however, occur in
the xeric sand areas of the Central Highlands of Flor-
ida".
Aside from the western distribution (Texas, Okla-
homa, New Mexico;fide Luginbill & Painter, 1953:51),
it is wingless and has limited dispersal powers. We
believe the specimen was mislabelled.

Phyllophaga dtssimilis (Chevrolat)

This Cuban species has been intercepted several
times at Miami, Florida: 6-VI-50, 11-VI-50, 24-VI-50,
all by J. E. Porter, on aircraft from Havana, Cuba
[INHS]; 5-VI-56,B. B. Sugarman, from Cuba [FSCA];
2-V-57, D. A. Miller, at Hibiscus esculentusLinnaeus,
interception No. M-10281, from Cuba [FSCA].
Chapin (1932:201-202, fig. 35) redescribed and
figured the species, recording it from the following
Provinces in Cuba: Santa Clara, Camaguey, and
Oriente. Other Provinces represented in the INHS
include: Havana, Pinar del Rio, and Matanzas. Box
(1953:9) listed it as apestof sugarcane in Cuba. There
is no evidence that it is established in Florida.

Phyllophaga drakii (Kirby)

Early in this study, we believed this species was
found in Florida. However, on further study, all
specimens proved to be eitherpostrema (fig. 44, 104,
164, 224, 257, 347, 350) or ulkei (fig. 56, 116, 176,
236,262,371, 374); two species with similar genitalia.
It is illustrated here (fig. 14, 74, 134, 194, 243, 283,
286) to aid in this distinction. *
Luginbill and Painter (1953:92-93) recorded it

from nearly all eastern states, except Florida. It is a
very large (av. 22mm long), dark species, often easy to
pick out from large collections of mixed species.

Phyllophagafraterna Harris

Several older lists (Blatchley, 1929; Young &
Thames, 1949) recorded this species for Florida. Because
of the similarity of several species in this group (e.g.,
curialis, floridana, foxii, incuria, mississippiensis,
paternoi, pseudofloridana), some of which were sepa-
rated only recently, these probably were misidenti-
fied. There appear to be no valid Florida records for
this species.
It is illustrated here (fig. 22, 82, 142, 202, 247,
300, 303) and distinguished from floridana (fig. 18,
78,138,198,244,293,296) andpseudofloridana (fig.
47, 107, 167, 227, 259, 353, 356). Sincefraterna is
recorded from nearly the entire eastern half of the
United States (Luginbill & Painter, 1953), additional
study is needed to clarify the species in this complex.

Phyllophaga glabricula (LeConte)

Although no specimens were seen from Florida in
the present survey, Luginbill and Painter (1953:60)
recorded it from "the prairie States of the United
States and also in Florida (fig. 54)." The other states
shown on their map are Colorado, Kansas, Nebraska,
Oklahoma, and Texas. Since we were unable to find
confirmation of this record 1) in the USNM (where
many of Luginbill & Painter's specimens reside) or 2)
in the extensive collection and records of M. W.
Sanderson at the INHS, and 3) this is a very unlikely
distribution to include Florida, we doubt this record.
It is possible that the "glabricula" was somehow
mistakenfor "glaberrima", a ubiquitousandcommon
species in Florida. The genitalia of glabricula are
quite distinct and were illustrated by Luginbill &
Painter (1953: pi. 53, fig. 6-11).

Phyllophaga hirtiventris (Hurn)

One specimen bf this species was intercepted at
Lake Buena Vista (Orange Co. Florida), 25-III-87, by
C. Phelps in a nursery, on Yucca recurvifolia Salis-
bury, from Houston, Texas.
It is treated and illustratedbyLuginbill and Painter
(1953:57-58, pl. 53, fig. 1-5) and was considered
common in some areas. However, no specimens were
recorded from Florida, and no others were found
during our survey. Fattig (1944:17) recorded a single

Table 6.
List of misspelled use of Florida Phyllophaga names.

Misspelling Correct name Author, date: page

austricolia austricola Fall, 1929a:110 (printer's error)
cupullfornis cupuliformis Young & Thames, 1949:127
debelis debills Young & Thames, 1949:127
duvalls duvalus Young & Thames, 1949:127
ephilids ephilida Hayes, 1925:118
ephilidia ephilida Dala Torre, 1912:188
foster forsterl Fattig, 1944: 22; Jaques, 1926: 338, 1927: 315, 1928: 304;
Owens, 1950:62-63, 83, 88; Travis, 1933: 397,1934:317
futllus futilis Ritcher, 1966:86. 87, 97
futulis futilis Jaques, 1926:338, 1927:315, 1928:304
herticula hirticula Jaques, 1926:338
illlcis flicis Hayes, 1925:15, 81, table 57, 59
implicata Implicita Hayes, 1925:13, 56, 65; Jaques, 1927:315
anchophora onchophora Fattig, 1944:31
prununcilina prununculina Cartwright, 1939:285
subpruninosa subprulnosa Ray, 1967:45
taxidi taxodli Loding, 1945:105
tristic tristls Fattig, 1944:29

Georgia specimen from Tate, 27-VI-1932. Except for
that record, it appears to be distributed primarily in the
central part of the United States (i.e., Arkansas, Indi-
ana, Iowa, Kansas, Kentucky, Louisiana, Mississippi,
Missouri, Nebraska, Oklahoma, and Texas). The male
and female genitalia are similar in general shape to
latifrons, but distinct in specific characters.

Phyllophaga hondura Saylor

Three specimens of this Central American species
were found at a nursery in Apopka (Orange Co.,
Florida), 13-V-85, P. Gibson, in unrooted cuttings of
devil's ivy or golden pothos (Epipremnum aureum
(Linden & Andre) Bunt.), shipped from Honduras.
One of these was still alive when it was received in
Gainesville for identification.
The species is known from Honduras, Belize
(British Honduras), and Costa Rica (Saylor, 1943). In
Honduras, King (1984) found it common in areas with
between 1000 and 2000mm annual rainfall (normally
with a 2 year life cycle); he illustrated the genitalia
(fig. 5, p.44). Although there is no evidence that it is es-
tablished in Florida, the increased flow of plants from

Central America in the nursery industry, and the inter-
ception mentioned above, indicate the probability of
introduction.

Phyllophaga hubbelli Cartwright

This species may occur in northern Florida. It was
described from 3 specimens (1 male, 2 females) taken
in: Alabama, Calhoun Co., Oxford, 1.5mi. S., 18-VII-
38, Hubbell and Friauf (misspelled Freauf in original
description) [USNM, 1 male]; Georgia, Summerville,
4-VIII-37, P. W. Fattig, on hickory andred oak [INHS,
2 females]. No additional specimens have been re-
ported for this rare species. The distribution suggests
that it may be found in the panhandle of Florida.

Phyllophaga lobata (Fall)

There is a single specimen in the FSCA with a very old
handwritten label, indicating "Jackson Co., Fla.",
probably from the University of Florida Agriculture
Experiment Station collection; undoubtedly misla-
belled. A rare species found only in Arizona. It is
treated and illustrated by Luginbill & Painter (1953:19,

pl. 21, fig. 1-5).

Phyllophaga lodingi Sanderson

A single specimen in the FSCA is from Alachua Co.,
17-1-36, with no collector or specific locality. The
label is similar to that used for student collections at
the University ofFlorida; undoubtedly mislabelled. A
rare mountain species from Madison Co., Alabama. It
is treated and illustrated by Luginbill & Painter (1953:82-
83, pl. 62, fig. 8-14).

Phyllophaga omani Sanderson

This rare species is a member ofthe subgenus Phytalus
and is similar in appearance to georgiana and gracilis.
It is known from 7 specimens: Alabama, Bumsville,
20-VII-30, P. W. Oman holotypee, female); Georgia,
Prattsburg, 24-VII-30, R. H. Beamer, at light (al-
lotype, male); Georgia, Thomasville, 28-V-38, W. H.
Thames, Jr., on crab apple (1 male) [INHS]; "North
America" (Saylor, 1939:162-163; pl. 9, fig. 2a-e; 1
male); Mississippi, Biloxi (Lago, 1980; 1 male); Aiken
Co., South Carolina (Riley, 1988; 1 male); and Louisi-
ana, Washington Parish, Lee Memorial Forest, near
Sheridan, 7-VIII-85, C. B. Barr, mercury vapor and
blacklight sheet (Riley, 1988; 1 male).
Nothing else appears to be known about the spe-
cies. The record from Thomasville, Georgia, suggests
that it may occur in Florida. It was treated and the
genitalia illustrated by Saylor (1939:162-163, fig. 2a-
d).

Phyllophaga onchophora Chapin

A single specimen of this Cuban species was
found at Savannah, Georgia, by P. W. Fattig, at light.
The date was recorded by Sanderson (1942:42) as
"May 2,1929", butFattig (1944:31) listed itas "May
21,1929" andmisspelledthenameas "anchophora".
The species was described from 5 specimens
from the following localities in Cuba: Paso Estancia;
Trinidad, Santa Clara; Santiago de las Vegas, Havana;
Perico, Matanzas. Since 2 other West Indian species
youngi Cartwright and bruneri Chapin) are estab-
lished in Miami, another (puberula) was collected
once in Miami, and onchophora was found in a Geor-
gia port, it eventually may be found in Florida. It was

treated and the genitalia illustrated by Chapin (1932:184,
fig. 18).

Phyllophaga portorlcensis Smyth

A single specimen of this Puerto Rican species was
takenatPalm Beach Co.,Florida, Delray Beach, 17-V-
79, K. Stolley, on Dracaena deremensis Engl., from
Puerto Rico. It was intercepted live in a shipment of
plants, but is not likely to be established in Florida.
Along with the similar Phyllophaga vandinei Smyth,
this is the most economic species of Phyllophaga on
sugarcane in Puerto Rico (Wolcott, 1948:255-262;
Box, 1953:9). It is another member of the subgenus
Cnematachis.

Phyllophaga rugosa (Melsheimer)

A single, badly damaged, male specimen has the
label data: Florida, Alachua Co., Gainesville, Chan-
tilly Acres, 11-VI-68, F. S. Blanton, mosquito light
trap. Although this is a common species in the north-
east and midwest, it is not recorded from the tier of
states north of Florida (Luginbill & Painter, 1953: fig.
76). Additionally, hundreds of light trap samples over
30 years in Gainesville, and many from Chantilly
Acres, have produced no further specimens.
Because of the above, and knowing that the
collector operated traps in New York also, this appears
to be a label error. It probably is not found in Florida,
but for aid in identification, refer to the genitalia
illustrations in Luginbill & Painter (1953: pl. 71, fig. 6-
12).

Phyllophaga submucida (LeConte)

The only published Floridarecord for this species
is Luginbill and Painter (1953:29, fig. 17). Their
distribution map is similar to that for glabricula (an-
other questionable Florida record); including Arkan-
sas, Kansas, Missouri, Oklahoma, and Texas, with no
records for the tier of states north of Florida. We were
unable to verify the source for any Florida specimens
(including pers. comm. with M. W. Sanderson), and
we do not believe it occurs here. The genitalia were
illustrated by Luginbill & Painter (1953: pl. 33, fig. 1-
6).

Table 7.
Annotated Alphabetical Checklist of names used for Florida Phyllophaga
(synonyms are italicized, valid names are bold face)

aemula (Horn) 1887b:271
alpine (Linell) 1896:726
anxia (LeConte) 1850:226
apicata Reinhard 1939:58
austricola Fall 19296:216
austricolia Fall 1929a:110

biimpressa (Smith) 1889a:97
boops (Horn) 1887b:284
brevicollis (Blanchard) 1850:132
bruneri Chapin 1932:203
burmeisteri (LeConte) 1856:242
carolina (Fall) 1912:43
cephalica (LeConte) 1856:245
cerasina (LeConte) 1856:241
ciliata (LeConte) 1856:253
clemens (Horn) 1887b:227
clypeata (Hor) 1887b:283
comans (Burmeister) 1855:358
comosa Davis 1920:337
crenulata (Froelich) 1792:94
cupuliformis Langston 1924:450
deanii Luginbill 1928:78
debilis (LeConte) 1856:262
decidua (LeConte) 1856:246
diffinis (Blanchard) 1850:138
dispar (Burmeister) 1855:361
dubia (Smith) 1888:183
duvalus Robinson 1938:110
elizoria Saylor 1937:321

elongata (Linell) 1896:725
ephilida (Say) 1825:196
fimbriata (Burmeister) 1855:326
floridana Robinson 1938:110
forbesi Glasgow 1916:378
forsteri (Burmeister) 1855:325
futilis (LeConte) 1850:226
foxii Davis 1920:334
georgicana (Gyllenhal) 1817:77
georgiana Schaeffer 1909:382

georgiana (Horn) 1885:122
gibbosa (Burmeister) 1855:324
glaberrima (Blanchard) 1850:136
gracilis (Burmeister) 1855:361
grandior (Linell) 1896:727
hirticula (Knoch) 1801:79
hornii (Smith) 1889a:95
howei Sanderson 1937a:17
ilicis (Knoch) 1801:75
implicit (Horn) 1887b:262
inana (LeConte) 1856:242

valid species
synonym of anxia (LeConte)
valid species
valid species [described as subspecies of tristis]
synonym of debilis (LeConte)
misspelling of austricola Fall in original description;
synonym of debilis (LeConte)
synonym of profunda (Blanchard)
synonym of dispar (Burmeister)
synonym of anxia (LeConte)
valid species (subgenus Cnemarachis)
synonym of ephilida (Say)
synonym of uniforms (Blanchard)
synonym of anxia (LeConte)
synonym of prununculina (Burmeister)
synonym of ilicis (Knoch)
valid species
valid species
synonym of diffinis (Blanchard)
synonym of hirticula (Knoch)
valid species
valid species [described as subspecies of micans (Knoch)]
synonym of snbpruinosa (Casey)
valid species
synonym of futills (LeConte)
valid species
valid species
synonym of anxia (LeConte)
synonym of schaefferi Saylor
valid species; new name for pygidialisSchaeffer 1906,
not Brenske 1892:190.
valid species
valid species
synonym of ilicis (Knoch)
valid species
valid species
valid species
valid species
valid species
synonym of crenulata (Froelich)
preoccupied; new name: schaefferi Saylor 1937 (not
Horn 1885:122)
valid species (not Schaeffer 1909) (subgenus Phytalus)
synonym of futilis (LeConte)
valid species
valid species
synonym of profunda (Blanchard)
valid species
valid species
synonym of clemens (Horn)
valid species
valid species
synonym of gracilis (Burmeister)

infidelis (Horn) 1887b:253
insperata (Smith) 1889a:93
integra (LeConte) 1856:258;
jonesi Sanderson 1939:5
knochii (Schoenherr & Gyllenhal) 1817:75
latifrons(LeConte) 1856:241
linelli Saylor 137:321
lota Luginbill 1928:87
luctuosa (Horn) 1887b:254
lugubris (LeConte) 1856:248
lutescens (LeConte) 1856:249
mariana Fall 1929a 111
micans (Knoch) 1801:77
minor Linell 1896:728
murrea Sanderson 1950:90
nova (Smith) 1889a:95
obsoleta (Blanchard) 1850:131

okeechobea Robinson 1948:33
ovalis Cartwright 1939:353
pagilis Saylor 1937:321

panorpa Sanderson 1950:91
parva (Linell) 1896:726

parvidens (LeConte) 1856:259
perlonga Davis 1920:329
pilosicollis (Knoch) 1801:85
politula (Horn) 1887b:248
porcine (Hentz) 1830:253
postrema (Horn) 1887b:233
profunda (Blanchard) 1850:132
prnunuculina (Burmeister) 1855:360
pseudofloridana Woodruff & Beck
puberula (DuVal) 1856:56
puncticollis Blanchard 1850:133
pygidialis Schaeffer 1906:257
quadrata (Smith) 1889a:94
quercus (Knoch) 1801:72
rufiola LeConte 1856:256
rugosioides (Linell) 1896:728
schaefferi Saylor 1937:321

semicribrata (LeConte) 1856:247
serricornis (LeConte) 1856:247
skelleyi Woodruff & Beck
sororia (LeConte) 1856:246
subpruinosa (Casey) 1884:38
taxodii Langston 1924:449
tecta Cartwright 1944:32
tristis (Fabricius) 1781:39
ulkei (Smith) 1889a:94
uniforms (Blanchard) 1850:133
uninotata (Walker) 1866:323
vanalleri (Schaeffer) 1927:215
volvula LeConte) 1856:235
yemasseei Cartwright 1944:30
young Cartwright 1935:102

Table 7 (cont.).

valid species
synonym of anxia (LeConte)
synonym of clypeata (Horn) (not Say 1835:180-181)
synonym (?) of ilicis (Knoch)
valid species
valid species
synonym of implicit (Horn)
valid species
valid species
synonym of forsteri (Burmeister)
synonym of forsteri (Burmeister)
valid species
valid species
synonym of implicita (Horn)
synonym of elongata (Linell)
synonym of forsteri (Burmeister)
valid species (includes vanalleri Schaeffer)(subgenus
Phytalus)
valid species
valid species
synonym of glaberrima (Blanchard); new name for
parva Linell 1896, not Brenske 1892:180
valid species
synonym of glaberrima (Blanchard) (not Brenske
1892:180)
valid species
valid species
synonym of tristis (Fabricius)
synonym of forsteri (Burmeister)
synonym of ilicis (Knoch)
valid species
valid species
valid species
new species
valid species
synonym of anxia Blanchard
synonym of elizoria Saylor (not Brenske 1892:190)
synonym of postrema (Horn)
valid species
synonym of diffinis (Blanchard)
synonym of luctuosa (Horn)
valid species; replacement name for georgiana Schaeffer
1909:382, not Horn 1885:122.
synonym of forsteri (Burmeister)
synonym of futilis (LeConte)
new species
synonym of diffinis (Blanchard)
valid species
valid species
valid species
valid species
valid species
valid species
synonym of anxia (LeConte)
synonym of obsoleta (Blanchard): new synonym
synonym of gracilis (Burmeister)
valid species
valid species (subgenus Cnemarachis)

Table 8.
Summary of
Taxonomic
Character
States

aemula
anxia
apicata
bruneri
clemens
clypeata
crenulata
cupuliformis
debilis
diffinis
dispar
elizoria
elongata
ephilida
floridana
forbesi
forsteri
foxii
futilis
georgiana
glaberrima
gracilis
hirticula
hornii
ilicis
implicita
infidelis

s a as

I I I iI I
0. .55

S 5 S S 8u u . S S S2 -E

X X X X X X 17.4 24.8
X X X X X X X 172 23.6
X X X x x X X 10 13
x x x x x x x 7.8 10.3
X X X X X X X 9:5 11
X X X x X X X 15.9 18
X X X X X X X 14.8 22.2
x x x x x x X 13.8 17.8
x X X x x x X 9.7 12.5
XX X X X X X 14 16.8
X X X x x X X 112.9
X X X X X X X 14.4 17.3
X X X X X X X 12.4 15.5
X X X x x x X 13 17.8
X X X X X X 166 20.6
x x x x x X 13.6 17.9
X xx X X X X 14.4 18.5
X X X X X X X 6.5 19
X X X X X X 14.2 17.9
X X X X X X 11.614.5
X X X X X X X 13.1 16.
X X X X X X10 13.2
XX XX X X X 16.2 20
SX X X x X 19 23.1
X X X X X X X X 18.3 23.9
X X X X X X X 14.6 18.5
X x XX X X 18.3 20.'

'Long = 1/2 or more length of upper spur.
2Reduced less than 1/2 length of upper spur.

a

a a

E U

s.I

u u

Table 8.
Summary of
Taxonomic
Character
States

knochii
latifrons
lota
luctuosa
mariana
obsoleta
okeechobea
ovalis
panorpa
parvidens
perlonga
postrema
profunda
prununculina
pseudofloridana
puberula
quercus
schaefferi
skelleyi
subpruinosa
taxodii
tecta
tristis
ulkei
uniforms
yemasseei
young

X X X X X X X 9.723.3
X X X X X X X 12.8 19.3
X X X X X X X 9.6 13.7
X X X X X X X 17.823.6
X X XX X X X X 19.8 24
X X X X X X x 13 17.5
X X X X X X x 13 16.2
X X X X X X X 18.421.6
X X X X X x X 16.1 19.1
XX X X X X X X 13.922.3
X X X X X X X 208 4..
X X X X X X X 18.6 23.7
X X X X X X X 18.725.2
X X X X X X x 14 18.5
X X X X X X X 14 18.3
X X X X X X X 14.5 16.7
X X X X X X X 123 16.9
X X X X X X 15.1 18
X X X X X X 15 19.4
X X X X X X 14.1 16.6
X X X X X X x 10.6 15.9
X X X X X X X 16.1 21.8
X X X X X X X 10 14
X X X X X X X 12.7 24
X X X X x X 13.717.7
X X X X X X X 10 10.9
X X X X X X X 15.4 17.5
'Long = 1/2 or more length of upper spur.
2Reduced = less than 1/2 length of upper spur.

Key to Adults of Florida Phyllophaga

1. Tarsal claws toothed (fig. 379, 381, 382, 383, 384) ................................................................... 2
1'. Tarsal claws cleft (fig. 380, 383); subgenus Phytalus .... ........................... ....... 3

2(1). Hind and middle tibiae with incomplete carina; lateral tibial margin with separate edges or well marked
spines (fig. 426-427); 3 introduced Cuban species, known only in Miami area; see male genitalia 186,
228, 239, 240 subgenus Cnemarachis ............................. .. ........................................................ 4
2'. Hind and middle tibiae with complete transverse or oblique carina at middle; lateral tibial margin without
edges or spines; subgenus Phyllophaga ......................................... ...... ......... ......... ...... ........... 6

3(1'). Tarsal cleft with terminal (outer) tooth longest (fig. 380); male genitalia: symmetrical, with 2 long
projections, not united at tip (fig. 24, 84,144, 204); female genitalia: pubic process bifurcate, deeply di-
vided (fig. 305, 308) ........................................... ......................................... georgiana Horn
3'. Tarsal cleft with lower (inner) tooth longest (fig. 383); male genitalia: symmetrical, the tip united, with-
outprojections (fig. 38,98,158,218); female genitalia: pubic process missing,tip ofplates notprolonged
or deeply divided (fig. 335, 338) .................................................... obsoleta (Blanchard)

4(2). Body with dark pattern (often metallic) on elytral suture, disc of pronotum (fig. 476); aedeagus with
group of spines pointing dorsally toward large paired hooks, covered by a dorsal shield (fig. 186); dis-
tribution: Cuba, Miami to Ft. Lauderdale and Naples; length 7.8-10.3mm .................. bruneri Chapin
4'. Body uniformly pale-colored, never dark on elytral suture or disc of pronotum; genitalia as in fig. 228,
239,240; length 14.5-17.5m m .......................................... ........................................................... 5

5(4'). Body glabrous, shiny, without dorsal pubescence; male genitalia with parameres projecting into boot-
shaped tip (fig. 239), aedeagus asymmetrical, flared at tip (fig. 59); female genitalia: fig. 377-378;
distribution: Bahamas, Miami (Brickell Hammock); length 15.4-17.5mm .............. young Cartwright
5'. Body pubescent with short hairs; parameres ofmalegenitalia without boot-shaped tip, smoothly rounded
to tip (fig. 228), aedeagus symmetrical to near the produced, nearly pointed tip (fig. 48); female geni-
talia: fig. 354, 357; distribution: Cuba, Miami (1 record); length 14.5-16.7mm ...... puberula (DuVal)

6(2'). Body above pubescent, hairs long or short .................................... ................................... 7
6'. Body above glabrous, if setae present, widely scattered .......................................................... .... 19

7(6). Setae of elytra arranged in rows; male genitalia asymmetrical (fig. 27), female genitalia: fig. 311, 314
........................................................ . ........................................................ hirticula (K noch)
7'. Setae of elytra variable but never in rows; male genitalia mostly symmetrical (except ilicis)............ 8

8(7). Male genitalia asymmetrical (fig. 29,89,149,209,250); female genitalia with elongate, bifurcatepubic
process (fig. 313, 316) ..................................................................................................... ........... ilicis
8'. Male genitalia symmetrical (fig. 1,4, 9, 15, 16, 36, 39, 41, 42, 55); female genitalia variable, but not
exactly as above .... ......................................... .................. ........................ ..... .................. 9

9(8'). Hairsofdorsum exceptionally long, especially on pronotum; small (10-14mm); clypeus entire (fig.421);
aedeagus protruding between parameres (fig. 4, 55); tristiss complex] .......................................... 10
9'. Hairs ofdorsum variable, from short-recumbent to long-erect; larger (12.4-24mm); clypeus emarginate;
aedeagus not protruding between parameres. ........................................ .................................... 11

10(9). Aedeagus with short terminal spinose process (fig. 184,468,471), area behind it shaped as wish-bone
(fig. 4, 64, 124, 470); females indistinguishable from tristis; in Florida, recorded only from Alachua &
Union Co ................................................................................................ .. .......... apicata Reinhard
10'. Aedeagus with long curved terminal spinose process (fig. 235, 469, 473), area behind not wish-bone
shaped (fig. 55,115,175,472); females indistinguishable from apicata; in Florida, recorded only from
panhandle ................................................................................................................. tristis Fabricius

11(9'). Larger species (19.8-24mm); dorsal hairs uniform, not longer on pronotum; male genitalia with trilobed
appearance (unique) (fig. 36, 96,156, 216); female genitalia: 329, 332 ........................ mariana Fall
11'. Size variable (12.4-24.8mm), butmostly smaller species (12.4-20mm); dorsal hairs variable, from short
to extremely long and recumbent on pronotum; male genitalia not trilobed; female genitalia variable
........... ...... .............................................................................. 12

12(11'). Pubescence variable, sometimes long on pronotum; lower male posterior tibial spurs both movable; body
shape widest behind, not parallel sided; male genitalia variable (fig. 1, 9,15, 39, 42, 51) .............. 14
12'. Pubescence short, not obvious; lower male posterior tibial spur fixed and reduced; body shape nearly
parallel sided; male genitalia with parameres produced, not united at tip (fig. 16, 41).................... 13

13(12). Smaller (12.4-15.5mm); tips of male parameres concave on inside (fig. 16, 498); tooth of male tarsal
claws basal, but noticeable; female genitalia: fig. 288, 291 ........................................ elongata Linell
13'. Larger(16.1-19.1mm); tips ofmaleparamres convex on inside (fig. 41,499); toothofmaletarsalclaws
missing; female genitalia: fig. 341, 344 ............................................................... panorpa Sanderson.

14(12). Male genitalia in caudal view with almost circular opening, the parameres united at tip (fig. 1, 9) ....
15...................................................................................................................................................... 515
14'. Male genitalia in caudal view with opening elongate, oval, the parameres elongate and somewhat
carinate (fig. 15, 39, 42, 51) .................................................................................................... 16

15(14). Male genitalia with parameres prolonged at tip, the notch between acute (fig. 1,61, 121, 181); female
genitalia (fig. 263,266) with outer angle of pubic process truncate, not pointed .......... aemula (Horn)
15'. Male genitalia with parameres not prolonged at tip, the notch between wide, obtuse (fig. 9,69,129,189);
female genitalia (fig. 275,278) with outer angle of pubic process produced, pointed ........................
................................... ................................................................ ........................ cre nulata (Froelich)

16(14'). Pronotal pubescence similar to that of elytra, short, recumbent; male genitalia deeply, broadly notched
above aedeagal opening, parameres divided for most of their length, the tips touching (fig. 42) .....
.......................... ............................................................................ ................ parvidens (LeC onte)
16'. Pronotal pubescence longer than that of elytra, long and some erect (fig. 586, 587); male genitalia not
notched at top of aedeagal opening, parameres divided for short distance, the tips diverging, pointed (fig.
15, 39, 5 1)..................................................................................................................................... 17

17(16). Male genitalia with double carina above aedeagal opening; outer margin of parameres indented opposite
where they unite below (fig. 15, 39) ....................................................................... ................ 18
17'. Male genitalia not carinate above aedeagal opening; outer margin of parameres evenly arcuate (not
indented) from top to tip (fig. 51, 111, 171, 231; female 360, 363) .................................. skelleyi n.sp

18(17). Male genitalia in ventral/caudal view with the carinae above aedeagal opening on each side of a deep
groove, sides more swollen (fig. 39, 99,159, 219,494); possibly diumal ........ okeechobea Robinson
18'. Male genitalia in ventral/caudal view with the carinae on each side of raised area, not deeply grooved,
sides less swollen (fig. 15, 75, 135, 195,495); nocturnal............................................ elizoria Saylor

19(6). Body above pruinose (velvety), sometimes slightly iridescent ...................................................... 20
19'. Body above mostly shining, never pruinose, not iridescent ............................................................ 24

20(19). Clypeus entire (fig. 423); 8th sternite of male with bifurcate process (fig. 542) (male genitalia: fig. 33,
93, 153,213; female genitalia: fig. 323, 326)............................................ latifrons (LeConte)
20'. Clypeus emarginate; 8th sternite of male without process; genitalia not as above ......................... 21

21(20). Lower spur of posterior tibia (male) missing (fig. 400); tooth of anterior tarsal claw postmedian; male
genitalia: fig. 46, 106, 166, 226; female genitalia: fig. 349, 352................ prununculina (Burmeister)
21'. Lower spur of posterior tibia (male) variable, but always present; tooth of anterior tarsal claw median;
male & female genitalia not as above ................................................. .... .................................. 22

22(21'). Lowerspurofposteriortibia(male) movable (fig. 401); body colorreddish tochestnut brown, neverdark
purplish; body parallel sided: male genitalparameres united: fig. 49,109,169,229; female genitalia: fig.
355, 358.................................................. ..................................................... quercus (Knoch)
22'. Lower spur of posterior tibia (male) fixed (fig. 399); body color purplish (at least the pruinose sheen);
body wider behind; male genitalia with parameres separated (fig. 10, 52) ................................... 23

23(22). Tooth of anterior tarsal claw strong, acute; male genitalia: fig. 52, 112, 172, 232; female genitalia: fig.
361, 364............................................................................................................. subpruinosa (Casey)
23'. Tooth of anterior tarsal claw weak, obtuse (fig. 382); male genitalia: 10, 70,130,190; female genitalia:
276, 279........................... ......................................................................... cupuliformis Langston

24(19). Male genitalia asymmetrical ..... .................................................. ......... ................................ 25
24'. M ale genitalia sym m etrical ......................................................... ......... ................................... 43

25(24). Antenna 9-segm ented. .................................................................................................................... 26
25'. Antenna 10-segm ented.................................................................................................................... 28

26(25). Small (9.7-12.5mm), pale yellow; tooth of anterior tarsal claw post median; male genitalia with
parameres overlapping at tip (fig. 11, 71,131,191); female genitalia with pubic process pointed, a deep
V-shape division at tip (fig. 277, 280) .......................................................................... debilis (Horn)
26'. Larger (14-18.5mm), dark brown; tooth of anterior tarsal claw median; male genitalia with parameres
not overlapping at tip (fig. 12, 30); female genitalia with pubic process truncate at tip (fig. 281,317).
27...................................................................................................................................................... 2727

27(26'). Male genitalia (fig. 12,72,132,192) with large cusps on each paramere, tips broad; female genitalia (fig.
281,284) with pubic process projecting laterally with about 13 long setae .......... diffnis (Blanchard)
27'. Male genitalia (fig. 30,90,150, 210,251) with parameres projecting to pointed tip, without large cusps
on each side; female genitalia (fig. 317,320) with pubic process projecting forward, truncate, with about
4 long setae .............................................................................................................. implicit (Horn)

28(25). Lower posterior tibial spur (male) movable; male genitalia (fig. 31, 50); female genitalia: 318, 359 ..
..................................................................................................................................................... 29
28'. Lower posterior tibial spur (male) fixed; genitalia variable, not as above.................................... 30

29(28). Male antenna club shorter than stem; body oval, shiny, convex, larger (18.3-20.7mm); male genitalia:
fig. 31, 91, 151,211,252; female genitalia: fig. 318, 321 ........................................ infidelis (Horn)
29'. Male antennal club equal to stem; body more parallel, more rugose, elytra more flattened, smaller (15.1-
18mm); male genitalia: fig. 50,110, 170, 230, 260; female genitalia: fig. 359, 362.. schaefferi Saylor

30(28). Male genitalia with nearly round opening to aedeagus, paramere tips overlapping below, pointing toward
each other (fig. 43, 44, 56) ............................................................................................................ 31
30'. Male genitalia with parameres elongate or with large cusps, the opening to aedeagus recessed more
deeply, variable in shape at the parameres which never overlap at tip (fig. 2, 18, 20,21, 28, 32, 35, 40,
45, 47, 54)...................................................................................................................................... 33

31(30). Male with right paramere produced outward, the bottom border truncate, broad, basal piece exception-
ally long (fig. 43,103,163,223,256); female genitalia: fig. 343,346; rare species in Florida panhandle
............................. .................................................................................................. p erlonga D avis
31'. Male genitalia nearly symmetrical except for overlapping paramere tips, right paramere similar to left,
not produced outward, basal piece normal length (fig. 44, 56); female genitalia: fig. 347, 371; more
common species, extending to peninsula.................................................................. ................ 32

32(31'). Male genitalia with right paramere having 2 posteriorly projecting processes (fig. 224); female genitalia:
fig. 347, 350 ...................................................................................... ............... postrema (Horn)
32'. Male genitalia with right paramere having 1 posteriorly projecting process (fig. 236); female genitalia:
fig. 371, 374; beetle: fig. 417 .................................................................................. ulkei (Smith)

33(30). Left paramere of male genitalia with diagonal carina on outside (fig. 258, 566); large species (18.7-
25.2mm); female genitalia: fig. 348, 351; Florida records only from Florida Caverns St Pk. ............
.............................. ....................................................................... ................ profunda (Blanchard)
33'. Left paramere of male genitalia with no carina on outside; size variable (14mm-23.6mm); female
genitalia not as above ................................................................................................................... 34

34(33). Parameres of male genitalia pointing downward, nearly pointed at tip, barely asymmetrical, no large
cusps formed inside parameres (fig. 2, 62, 122, 182); female genitalia with truncate, bifurcate pubic
process (fig. 264,267); Florida records only from Torreya St. Pk. .......................... anxia (LeConte)
34'. Parameres of male genitalia usually not projecting downward, but rounded parameres forming large
cusps (fig. 18,20,21, 28,32, 35,40,47, 54); female genitalia not as above; some widespread common
species .......................................................................................................................................... 35

35(34). Parameres of male genitalia in ventral view (fig. 100) pointing in opposite directions, posteriorly (left)
and anteriorly (right), many long setae on inside edge of parameres; female genitalia with long internal
process and broadly divided pubic process (fig. 337); rare, large (18.4-21.6mm), oval, shiny species,
known only in western Florida panhandle; beetle: fig. 416 ...................................... ovalis Cartwright
35'. Parameres of male genitalia in ventral view not pointing in opposite directions, with only a few setae on
inside; female genitalia not as above; size and shape variable ...................................................... 36

36(35). Male antennal club shorter than stem; large (17.8-23.6mm),oval, piceous species with coarsely punctate
pronotum; male genitalia: fig. 35, 95, 155,215,254; female genitalia: fig. 325, 328 luctuosa (Horn)
36'. Male antenna club equal in length to stem; size, shape, and color variable; male and female genitalia not
as abo ve ....................................................................................................................................... 37

37(36). Right internal process of male paramere modified into a hook or S-shape, the leftparamere with a double
cusp, formed by a carina medially (fig. 28,88,148,208,249,529); female genitalia: fig. 312,315; Florida
panhandle .................................................................................................................... hornii (Sm ith)
37'. Right internal process of male paramere not modified into a hook, the left paramere with only 1 cusp,
no median carina; female genitalia not as above ..................................................... ................. 38

82

38(37). Left paramere with posteriorly projecting tooth at top (fig. 200), top ofphallobase with medial knob (fig.
20); female genitalia: fig. 295, 298; small (14.4-18.5mm), oval, castaneous, shiny ....................
......................................................................................................................... forsteri (Burm sister)
38'. Leftparamere notprojecting posteriorly into a tooth; top ofphallobase withoutnoticeable knob; genitalia
not as above; size variable (14-23.6mm), shape and color variable ........................................ 39

39(38). Pronotum coarsely punctate, the median line noticeable; elytra castaneous with head and pronotum
darker; large (19.7-23.3mm); male genitalia: fig. 32, 92,152, 212; 253; female genitalia: 319, 322 ..
................................................................................................................... knochii Schoen. & G yll.
39'. Pronotum not coarsely punctate, the median line not obvious; color nearly uniform; size variable (14-
21.8mm); male genitalia: fig. 18, 21,47, 54; female genitalia: fig. 293, 299,353,366 .................. 40

40(39'). Color reddish brown to castaneous; large (16.1-21.8mm), oval, shiny, striae obliterated; male genitalia:
fig. 54, 114, 174, 234, 261; female genitalia: fig. 366, 369; common at Gainesville ...................
......... ................................. .............................................................................. .. tec ta C artw right
40'. Color dark brown to black; size variable (14-20.6mm), shape variable, elytra less shiny because of strial
indications and punctures; genitalia not as above (male: fig. 18, 21,47; female: 293,299, 353);fraterna
com plex ........................................................................................... .......... ................ 41

41(40). Internal process of right paramere sharply pointed, male genitalia: fig. 21, 81, 141, 201, 246; female
genitalia: 299, 302 [Sanford, Florida]; male: fig. 510-514; female: 515,516 [South Carolina], Florida
record questionable ................................................ ........................................... foxii Davis
41'. Internal process of right paramere not sharply pointed, flat and broad, male genitalia: fig. 18,47; female
genitalia: fig. 293, 353 ......................................................... .................................... 42

42(41'). Size larger (16.6-20.6mm); internal process of right paramere directed posteriorly (tip often hidden by
carina or cusp of right paramere), the area above aedeagal opening not swollen into distinct knob from
right lateral view; male genitalia: fig. 18,78,138,198,244; female genitalia: fig. 293,296; peninsular
Florida (see map fig. 504) ............................................................................. floridana Robinson
42'. Size smaller (14-18.3mm); internal process of right paramere directed upward (tip usually visible from
right lateral view), the area above aedeagal opening swollen into a distinct knob (fig. 571, 572); male
genitalia: fig. 47,107,167,227, 259; female genitalia: fig. 353,367, 573; north Florida, south Georgia
....................................................................................................................... pseudofl oridana n. sp.

43(24). Antennae 9-segmented; clypeus entire; mostly small species (9.5-15.9mm), except clypeata (15.7-
18mm); male genitalia: fig. 7, 8, 13, 34, 53, 58; female genitalia: fig. 270,271, 282, 324,365,373 ..
.................................................... ............................................................................................... 44
43'. Antennae 10-segmented; clypeus emarginate; some larger species (13-17.9mm), except gracilis (10-
13.2mm); male genitalia: fig. 17,19,23,25,26,57; female genitalia: fig. 289, 294, 301,306,307,372.
................................................ ................................................................................................ 49

44(43). Lower male posterior tibial spur movable; male genitalia with parameres long, cylindrical (fig. 188,
193); female genitalia with pubic process long, cylindrical (fig. 271,282)...................................... 45
44'. Lower male posterior tibial spur fixed; male genitalia variable, but not long, cylindrical (fig. 187,214,
233, 238); female genitalia variable (fig. 270, 324, 365, 373) ...................................................... 46

45(44). Larger (15.7-18mm); color castaneous to brown; male genitalia: fig. 8,68, 128, 188; female genitalia:
fig. 271, 274 .................................... ... ................................................................ clypeata (Horn)
45'. Smaller (11-12.9mm); color dark brown to piceous; male genitalia: fig. 13, 73, 133, 193; female
genitalia: fig. 282, 285 ...................................................... ................................ dispar (Burm sister)

46(44). Anterior tarsal claw of male with tooth median: male genital parameres sharply pointed at tip, with a knob
above directed inward (fig. 34,94,154,214); female genitalia unique with 3 lobes on pubic process, the
median directed upward and state (fig. 323, 327) ..................................... .... ota Luginbill
46'. Anterior tarsal claw of male with tooth postmedian (nearer base than tip); male genitalia variable (fig.
7, 53, 58), but parameres not pointed; female genitalia not trilobed (fig. 270, 365,373) ................ 47

47(46'). Larger (10.6-15.9mm); male genital parameres flattened in caudal view, indented laterally at above half
their length (fig. 53,113,173,233); pubicprocess of female genitalia bifurcate, the tips pointed(fig. 365,
368); specific to cypress ..................................... .............................................. taxodii Langston
47'. Smaller (9.5-11mm); male genital parameres not flattened in caudal view, not indented laterally (fig. 7,
58); pubic process of female genitalia either not bifurcate, or tips truncate (fig. 270, 373) ............ 48

48(47). Male genital parameres more than twice as long as wide, with cluster of long setae internally and on tips,
and no secondary process on caudal face of parameres (fig. 7, 67, 127, 187); pubic process of female
genitalia not bifurcate, tip triangular (fig. 270, 272) ...................................... clemens (Horn)
48'. Male genital parameres about as wide as long, without setae between or at tips, and an extra process on
caudal face of parameres (fig. 58, 118, 178, 238); pubic process of female genitalia bilobed, the tips
truncate (fig. 373, 376) ............................................................................... yemasseei Cartwright

49(43). Smaller (10-13.2mm); male genital parameres prolonged downward, pointed, sickle-shaped from side
view, setae projecting posteriorly along inside of curve (fig. 26,86,146,206); female genitalia: fig. 307,
310 .................................................................................................................... gracilis (B rm sister)
49'. Larger (13.1-17.9mm); male genital parameres not as above (fig. 17,19, 23, 25, 57); female genitalia:
fig. 289, 294, 301, 306, 372 ..................................................................... ................................. 50

50(49). Lower male posterior tibial spur reduced (less than half length of upper movable spur); male genitalia:
fig. 17, 25, 57; female genitalia: fig. 289, 306, 372........................................................................ 51
50'. Lower male posterior tibial spur long (more than half length of upper movable spur); male genitalia: fig.
19, 23; female genitalia: fig. 294, 301 ........................................ ........... .................................. 53

51(50). Male genitaliaelongate, cylindrical, parameres divided, resembling 4 long fingers, theouter(lower) ones
longerand strongest; malegenitalia: fig. 17,77,137,197; female genitalia: fig. 289,292; Florida records
doubtful ........................................................................................................... ........... ephilida (Say)
51'. Male genitaliaelongate or not,parameres notproduced into finger-like projections; male genitalia: fig.
25, 57; female genitalia: fig. 306, 372; common Florida species .................................................. 52

52(51'). Male anterior tarsal claws with tooth median; male genitalia 3 times longer than high (fig. 57,117,177,
237); pubic process of female genitalia long, cylindrical, notbifurcate at tip, no setae on tip (fig. 372,375)
....................................................................................................................... uniform is (B lanchard)
52'. Male anterior tarsal claws with tooth post median (nearer base than tip); male genitalia about 2 times
longer than high (fig. 25, 85, 145,205); pubic process of female genitalia flattened ventrally, bifurcate,
the tips with setae (fig. 306, 309) ................................................................. glaberrima (Blanchard)

53(50). Male genital parameres forming a cup-shape below, without inward projecting fingers at caudal apex or
internally (fig. 28, 83,143,203); pubic process of female genitalia broad, resembling superior plates, not
cylindrical (fig. 301,304); male anterior tarsal claws with median tooth; male lower posterior tibial spur
fixed, modified into unique, long, boot-shaped process (fig. 395) ............................ futilis (LeConte)
53'. Male genital parameres forming a cylinder, not cup-shaped below, but with caudal and ventral (median)
finger-like projections (fig. 19, 79, 139, 199); pubic process of female genitalia prolonged into a
cylindrical shape (fig. 294,297); male anterior tarsal claws with tooth postmedian; male lower posterior
tibial spur fixed, half length of upper .................................................................. forbesi Glasgow

Phyllophaga aemula (Horn)
(fig. 1, 61, 121, 181,263,266, 464,465)

Lachnosterna aemula Horn 1887b:271.
Phyllophaga aemula, Glasgow 1916:373.

TYPE LOCALITY: "northern Georgia".

DIAGNOSIS: Superficially it is similar to the follow-
ing hirsute Florida species (see genitalia figures in
parentheses for comparison): crenulata (male: 9, 69,
129,189; female: 275,278); elizoria (male: 15,75,135,
195; female: 287,290); ilicis (male: 29, 89, 149, 209,
250; female: 313, 316); mariana (male: 36, 96, 156,
216; female: 329,332); parvidens (male: 42, 102, 162,
222; female: 342, 345); and skelleyi (male: 51, 111,
171, 131; female: 360, 363).
In addition, aemula and parvidens have pruinescent
elytra under the pubescence, whereas crenulata does
not. In male ilicis the body is narrower, the antennal
club is longer, and the lower tibial spur is fixed. The
male genitalia of aemula have the bottom of the claspers
projecting, but in crenulata they are gently rounded.

DESCRIPTION: Length: 17.4-24.8mm; Width: 9.6-
13.2mm. Shape: oblong, oval, robust. Color: brown.
Vestiture: pubescent, clothed with fine, short, recum-
bent hairs; surface pruinose. Antenna: 10-segmented;
.male club two-thirds length of stem. Clypeus: emargi-
nate; border narrowly reflexed. Tarsal Claws: tooth
large, acute, median. Male Posterior Tibial Spurs:
lower movable, two-thirds length of upper; both spurs
broad and attenuate. Female Genitalia: fig. 263 (ven-
tral), 266 (lateral). Male Genitalia: fig. 1 (caudal), 61
(ventral), 121 (dorsal), 181 (right lateral).

TAXONOMIC NOTES: A fairly uniform species,
with no synonyms having been created. Variation was
noted primarily in body color (from light to dark or
reddish brown). As with crenulata, the pubescence
sometimes appears more noticeable, due to the thick-
ness of individual setae and partial removal from wear.
The holotype male (Type No. 3683) and paratype
female (No. 3683), labelled "Ga.", from the Horn
collection, were examined at the MCZC; both have the
genitalia dissected. Also in the Horn collection are: (2)
"Haulover, Fla., March, 364", and (1) "Horn Coll. H
10259."

U.S. DISTRIBUTION (fig. 465): Luginbill & Painter
(1953: fig. 11) recorded it from Alabanma, Arkansas,

Florida, Georgia, Mississippi, New Jersey, North
Carolina, Oklahoma, South Carolina, and Virginia.
In addition, Reinhard (1950:37) recorded it from Texas
(Brazos Co.), and Riley (1988: map 2) recorded it from
Louisiana (8 localities in 7 parishes).
The Arkansas record was based on 1 specimen
from Crawford Co. (Sanderson, 1944:21). Smith
(1910:319) saw only 1 from New Jersey (DaCosta).
Loding (1945:105) listed 8 counties in Alabama, and
Langston (1927:63) listed 9 records from Mississippi.
Cartwright (1934:268) saw a single specimen from
Clemson, South Carolina. Although the type locality
is "northern Georgia," Fattig (1944:27) saw only 5
specimens in his extensive Georgia survey (Albany,
Head River, Thomasville).

9 1

FLORIDA DISTRIBUTION (fig. 464): Our south-
ernmost record is Ocala (Marion Co.), and it probably
occurs from there north and west to include the entire
panhandle. Smith (1889b:518) reported specimens
from Haulover, Florida collected by Schwarz, and
these are the basis for Blatchley's (1929:69) record
from "Haulover (Sz. Ms. and Fall)". He apparently
saw no Florida specimens. We believe these specimens
may be a misidentification and represent one of the
elizoria complex. Young and Thames (1949:128)

dO""'

repeated Blatchley's record and added: "locally common
in northern part of state."

BIOLOGY & ECOLOGY: Luginbill & Painter
(1953:20) listed the adult season as May to July. Our
earliest Florida record was April 18 (Florida Caverns
State Park) and the latest was October (Gainesville). At
Gainesville it was most abundant in July and August.
There is a curious note on behavior by Schwarz
(1891:241) which follows: "In 1875 while in camp at
Haulover Canal, in Florida, I had occasion to observe
Lachnosterna aemula, which in the earlier part of
March was flying about after dark, but also in the early
morning, when it was still so dark that the flying beetles
could only be heard but not seen."
Smith (1889b:518) erroneously cited this reference
to read "... Haulover, Fla., March 11-13 ... they fly
shortly before sunrise, instead of at dusk and early
evening, as do most of the other species." [italics ours].
The identity of Schwarz's specimens is in doubt, since
we have not found aemula that far south, and in 1891
the 2 species that we suspect he had (elizoria or oke-
echobea) were not yet described. However, the reference
to a species having early morning flight gave us another
clue to the habits of the rare okeechobea (see discussion
under that species).

Adult Host Plants: Beech, ebony, pine, rose, willow
[families] (Luginbill & Painter, 1953:20); cypress, pine
(Langston, 1927b:63); willow oak (Sanderson, 1944:21);
red oak, persimmon (Fattig, 1944:27); pecan, grape
(Reinhard, 1950:47); dogwood (Riley, 1988:72).

Immatures: Only the first instar larva has been de-
scribed, and care should be used when comparing it
with the third instar of other species (e.g., in our Key).
On larval characters, Boving (1942:29-30, fig. 2, 8)
placed it in his group 1, along with crenulata, rubigi-
nosa, and parvidens. His description follows: "Poste-
rior part of labrum with no setae. Anterior marginal
region of frons with 2 setae on each side. Epicranium
on each side opposite concave posterior part of frontal
suture and epicranial suture with 2 setae. Dorso-molar
region of right mandible (Fig. 4) with an oblique series
of about 5 setae at the anterior part of mola and 15 to 20
setae in a patch behind; dorso-exterior region with a
few punctures or usually bare; scrobis with a longitudi-
nal row of 6 punctures; ventro-lateral carina without
setae. Maxillary articulating area ventrally with about
5 short, thick, dark granules (Figs. 2, 6). Epipharynx
with 5 large heli in a distal row and 2 short ones in a
proximal row. Proplegmatium (Fig. 5) distinct, subel-

liptical, moderately wide with 7 to 10 preplegmata;
chaetoparia with few, usually setula-bearing punc-
tures; crepidal punctures about 4. Raster (Fig. 8) with
inversely spatulate septula; each palidium with one
irregular row of from 25 to 27 straight, sharply pointed,
moderately long, densely set pali; preseptular setae
about 12, long, and arranged in three very irregular
transverse rows. Hatching spine (Fig. 3) represented by
a weakly sclerotized, thin, yellowish plate."

SPECIMENS EXAMINED: 450 from the following
9 Florida counties: Alachua, Gadsden, Jackson, Jeffer-
son, Leon, Liberty, Marion, Santa Rosa, and Wakulla.
Over 255 specimens, representing 111 records, were
from Gainesville. For complete data, see Appendix 1.

SELECTED REFERENCES: Blatchley, 1929:69; Boving,
1937:5; 1942:12-13,15,21,23,29-30,58, 61,62, figure 2-8,204-205;
Cartwright, 1934:268; Dalla Torre, 1912:183; Fattig, 1944:7-8, 27;
Glasgow, 1916:373-374; Horn, 1887b:265, 270-271,295; Langston,
1927b:62-63, 87, plate 10, figure 39; Leng, 1920:256; Loding,
1945:105; Luginbill & Painter, 1953:5, 20, figure 11, plate 22(1-4);
Reinhard, 1946:479; 1950:47; Riley, 1988:49,51,54,70-73,112,172,
208, figure 57-60, map 2, table 3-4; Sanderson, 1944:16, 21, table 1;
Schwarz, 1891:241; Smith, 1889a:518, figure 66, plate 58; 1910:319;
Travis, 1933:397; Woodruff, 1973:28; Young & Thames, 1949:128.

Phyllophaga anxia (LeConte)
(fig. 2, 3, 62, 63,122, 123, 182, 183, 241, 242, 264,
267,466,467)

Lachnosterna anxia LeConte 1850:226.
Ancylonycha brevicollis Blanchard 1850:132.
Ancylonycha puncticollis Blanchard 1850:133.
Lachnosterna cephalica LeConte 1856:245.
Ancylonycha uninotata Walker 1866:323.
Lachnosterna dubia Smith 1888:183.
Lachnosterna insperata Smith 1889a:93.
Lachnosterna alpina Linell 1896:726.
Phyllophaga anxia, Glasgow 1916:371.

TYPE LOCALITY: no specific locality, but pre-
sumed to be "Lake Superior".

DIAGNOSIS: Superficially itis similar to several dark
glabrous Florida species, but is easily distinguished by
the genitalia. As a member of thefusca group, the male
genitalia are asymmetrical (but only mildly) (fig. 2,62,
122, 182, 241), and the female genitalia (fig. 264,267)
have a distinct transverse fold on the basal plates. Of
Florida species, genitalia are similar only to implicita
(male: 30,90, 150,210,251; female: 317,320). However,

that species has 9-segmented rather than 10-segmented
antennae.

DESCRIPTION: Length: 17.2-22.5mm (23.6mm,
Riley); Width: 9.3-11.6mm (12.6mm, Riley). Shape:
oblong, ovate, subdepressed. Color: dark brown to
black. Vestiture: glabrous, moderately polished. An-
tenna: 10-segmented; male club length nearly equal to
stem. Clypeus: emarginate; border moderately reflexed.
Tarsal Claws: tooth strong, median. Male Posterior
Tibial Spurs: lower fixed, half length of upper. Female
Genitalia: fig. 264 (ventral), 267 (lateral). Male Geni-
talia: southern form: fig. 2 (caudal), 62 (ventral), 122
(dorsal), 182 (right lateral), 241 (left lateral); northern
form: fig. 3 (caudal), 63 (ventral), 123 (dorsal), 183
(right lateral), 242 (left lateral).

TAXONOMIC NOTES: With the 7 synonyms listed
above, it is obvious that there has been much confusion
about anxia. Variation in body size and in genitalia are
noticeable. Two distinct types of male genitalia (north-
ern and southern forms) are recognized (fig. 2,3,62,63,
122, 123, 182, 183, 241,242). Our few Florida speci-
mens are clearly the southern form. It has the widest
distribution of any U.S. species. Luginbill & Painter
(1953:80) mentioned that Utah specimens are half the
size of those from other parts of the country.

Space does not permit a thorough accounting of the
above synonymy, which was established by Glasgow
(1916:371). This is one of the species which needs
more detailed study, especially the internal sac, to
clarify the extensive variation.

U.S. DISTRIBUTION (fig. 467): Luginbill & Painter
(1953: fig. 73-74) recorded it from every state except
Arizona, California, Florida, Nevada, West Vir-
ginia and Wyoming. They also list 11 provinces in
Canada, making this the widest distributed of all U.S.
Phyllophaga.

FLORIDA DISTRIBUTION (fig. 466): The 4 Flor-
ida specimens were all taken at Torreya State Park
(Liberty Co.), the source for several records of northern
species in this relictual area along the Apalachicola
River. These are the first Florida records.

BIOLOGY & ECOLOGY: Because there is a north-
ern and southern form, based on genitalic differences,
some of the references to biology may refer to one or the
other. Our 3 Florida records are for April, May, and
July.
Forbes (1916:227-228) called it a distinctly northern
species in Illinois where it apparently has a 3-y;ear
cycle, collected as early as April 15 and as late as July
8. In North Carolina, Brimley (1938:203) reported
collecting it in the ground or under stones or logs in
winter and early spring. Hayes (1929:66) cited a report
that it had a 4-year life cycle in Manitoba, Canada.
Because of its economic importance and its widespread
distribution, there havebeen more studies (and resulting
publications) on this species than any other in the U.S.
Space permits only citing the references by subject
matter as follows:
Tiphiidae (Hymenoptera) parasites (Berberet &
Helms, 1970; Lim, Stewart, & Yule, 1981; Rivers,
Mayo, & Helms, 1979); gregarine parasites (Berberet
& Helms, 1969); mites (Jarvis, 1964, 1966; Oseto &
Mayo, 1975); anatomy and histology of adults and
larvae (Berberet & Helms, 1972); life history and
control in Canada (Hammond, 1948); general natural
enemies (Lim, 1979; Lim, Stewart, & Yule, 1981; Lim,
Yule, Stewart, 1980; Petch & Hammond, 1925, 1926);
oviposition (Sweetman, 1927).

Adult Host Plants: Basswood, birch, maple, olive,
beech, bignonia, buckeye, dogwood, ebony, elm, heath,
honeysuckle, magnolia, mallow, pulse, rose, sumac,
sweet gale, tupelo, walnut, willow, laurel, maple, oleas-
ter, polemonium, witchhazel, goosefoot [families]

(Luginbill & Painter, 1953:80-81); nectarine, persim-
mon (Reinhard, 1950:45); ash, willow, persimmon,
oak, sumac, blackberry (Ritcher, 1940:109); elm, wil-
low, poplar, apple, cherry, boxelder, hackberry, linden,
mountain ash (Forbes, 1916:228, "preference for elm
and willow"); poplar, pecan, willow (Langston,
1927b:44); farkleberry, Chickasaw plum, grape, sweet
gum, smooth alder, wax myrtle, water oak, black jack
oak, sand blackberry (Luginbill, 1928:72); preferred:
willow, ash, elm, lilac; others: plum (blossoms and
leaves), apple (blossoms and leaves), currant, choke-
cherry, hazelnut, spirea, raspberry, gooseberry, dock,
nettle, artichoke, giant ragweed, strawberry, golden-
rod, peony (Sweetman, 1931: table 19); quaking aspen,
large-toothed aspen, privet (Travis, 1934:329).

Immatures: The third instar larva was described by
Boving (1942:44, fig. 114-116), who places it in his
group 16, along withfusca, vehemens,fervida, margi-
nalis, and drakii (based on larval characters). His
description follows: "Posterior part of labrum with a
tranverse series of 5 to 7 long setae on each side.
Anterior marginal region of frons with a transverse,
somewhat irregular series of 5 to 7 setae on each side.
Epicranium on each side opposite posterior concave
part of frontal suture and epicranial suture with 3 or 4
setae. Dorso-molar region of right mandible (Fig. 116)
with a patch of about 30 setae; dorso-exterior region
with either from 3 to about 10, or from about 20 to 25
punctures; scrobis with about 7 punctures in an irregu-
lar, longitudinal row and occasionally with a single or
2 to 3 setae; ventro-lateral carina with 6 to 8 setae; baso-
lateral region with a patch of about 8 setae and some
punctures. Epipharynx with about 12 heli; propleg-
matium rather broad, long and spatulate with 11 to 15
curved proplegmata; right chaetoparia with numerous
punctures among the setae; crepidal punctures about
25. Raster (Fig. 115) with anterior third of septula oval,
tapering posteriorly into a subrectangular part; palid-
ium with one very irregular, in places double row of pali
numbering from 20 to 30 or more; palus (Fig. 114)
compressed, with concave sides and hooked at the tip;
majority of pali separated by a distance half as long as
a palus or shorter; preseptular setae 6 or more. (Length
of body about 40 mm.; width of head about 5.5 mm.)."
Ritcher (1966:87-88) included it in his key, but
included no illustrations.

SPECIMENS EXAMINED: several thousand, of
which only 4 were from Florida with the following
data: (1) Liberty Co., Torreya St.Pk., 14-IV-79, L. R.
Davis, Jr., at blacklight; (2) loc. cit., 11-VII-81, P. M.

Choate, Jr., on trees at night; (1) loc. cit, 21-22-V-83,
K. W. Vick, blacklight trap.

SELECTED REFERENCES: Berberet & Helms, 1969:395-
396; 1970:471-472, figure 1-3; 1972:1026-1027, 1031-1032, 1036,
1038, 1040-1042, 1044, 1047, 1050, 1052. figure 1-64. plate 1-7;
Blatchley, 1910:970, figure 390; Boving, 1937:3; 1942:6-7. 12, 23,
37, 45, 59, 61, figure 114-116, 237; Brimley, 1938:203; Cartwright,
1934:240; Chamberlin & Seaton, 1941:467, table 1; 1941:467, table 1;
Chamberlin, Fluke, & Callenbach, 1943:677-678, table 1-2; Cham-
berlin, et al., 1938:228, 230, 233,236, 238, table 1-2, 4-8; 1939:105,
table 1; Chandler, Taylor, & Deay, 1956:187; Criddle, 1918:3-4;
Crotch, 1874:60; Dalla Torre, 1912:190; Davis, 1918:4, 7, figure 2;
1919:81,84,106, 112; Dawson, 1922:215-216,221; Fattig, 1944:7-8,
20-21; Forbes, 1916:217-218,223-224,227-228,230,235,238-239,
241-248 252, 255; Glasgow, 1916:371, 374; Hammond, 1948:403-
416; Hayes, 1925:41,81; 1929:66; Henry & Heit, 1940:280-282; p, 1-
4; Hom, 1887b:294; Hudson, 1919:81-82 (as dubia); Jaques,
1926:338; 1927:315; Jarvis, 1964:207-210; 1966:401-409; Knaus,
1897:216; Langston, 1927b:34, 43, 83, plate 6, figure 24; LeConte,
1850:226; 1856:245; Leonard, 1926:424; Lim, 1979:i-xxi, 1-230,
illustr.; Lim, Yule, & Stewart, 1980:219-220;Loding, 1945:104;
Luginbill, 1928:56,71-72, figure 19, male A-E, female F-G; Luginbill
& Painter, 1953:10,79, figure73,plate69(1-12); McColloch & Hayes,
1923:30; McLeod & Schultz, 1988:95; Melsheimer, 1853:59; Naim &
Wong, 1965:33-34; Owens, 1950:33; Petch & Hammond, 1925:24-
28; 1926:85-91; Reinhard, 1950:45; Riley, 1988:70-74, fig. 3, map 3,
table 1,3-4; Ritcher, 1939:64; 1940:75,82-84, 86,109,128, figure 12,
plate 3; 1949a:19,25; 1966:86-87; Rivers, 1977:2-6,14-15,18,21,23,
27-31; Rivers, Mayo, & Helms, 1979:362-363, 372; Sanderson,
1944:16, 21, table 1; Schwardt, 1943:117; Shenefelt & Simkover,
1951:223; Sim, 1928:29-30,55, plate 7; Smith, 1889b:503; 1910:319
(as dubia); 1960:77; Sweetman, 1927:783,785-788, 790; 1931:401,
407-408, 411, 413-414, 416, 418-421, figure 4, table 10, 14, 17-19;
Travis, 1933: table 4; 1934:317, 328, 354, figure 16, plate 4, table 1;
Uhler, 1941:1-2, 8,10,12,14,16-17,19,21, figure 1,9,18,27,36-37,
plate 1,3,5,7,9; Wickham, 1894:223 (as dubia); Yeager, 1950:172,
176.

Phyllophaga apicata Reinhard
(fig. 4, 5, 64, 65, 124, 125, 184, 185, 265, 268, 403,
468,470,471,474,475)

Phyllophaga tristis apicata Reinhard 1939:58-59.
Phyllophaga apicata, Sanderson 1944:20.

TYPE LOCALITY: "College Station, Texas".

DIAGNOSIS: This and tristis are the only small (10-
13mm long), yellowish brown, hairy species in Florida.
These two are difficult to distinguish except by the
aedeagus (internal sac) of the males (females can be
identified only by association). In apicata (fig. 4, 64,
124, 184) there are 2 apical aedeagal processes; the
dorsal toothed near the apex, the distal process single
bladed, looped downward, then curved upward and
with apex entire.

~rinw ~

U

fig. 473. Phyllophaga tristis: aedeagus (right lateral view) (5mm
= 0.05mm).

DESCRIPTION: Length: 10.0-13.0mm; Width: 5.2-
6.5mm (7.3mm, Riley). Shape: oblong, slightly wider
behind. Color: rufotestaceous. Vestiture: pubescence,
long, erect on head and pronotum, shorter, suberect on
elytra; surface more shining than in tristis. Antenna:

10-segmented; male club shorter than stem. Clypeus:
entire, rounded to nearly straight medially; margin
broad, strongly reflexed. Tarsal Claws: tooth small,
antemedian, acute. Male Posterior Tibial Spurs: lower
movable, slender, obtuse, longer than first tarsal segment.
Abdomen: fig. 403 (venter). Female Genitalia: fig.
265 (ventral), 268 (lateral). Male Genitalia: fig. 4, 5
(caudal), 64, 65 (ventral), 124, 125 (dorsal), 184, 185
(lateral).

TAXONOMIC NOTES: This form was originally de-
scribed as a subspecies of tristis, along with amplicor-
nis and suttonana. These are now considered species,
although they are similar and confused in all literature
before 1939. The tristis complex has an extremely
broad geographic range and extensive external vari-
ation in vestiture. The only secure characters for
distinguishing the forms are in the aedeagus (internal
sac) of the male genitalia (fig. 468,470,471, vs. 469,
472,473). Females of all forms are presently insepa-
rable. The complex needs a thorough evaluation from
its entire range and an examination of the Fabrician
type of tristis.

U.S. DISTRIBUTION (fig. 475): Reinhard (1939:59)
listed paratypes from: Alabama (Mobile, Auburn);
Arkansas (Cody's Gap); Georgia (Zebulon, Perry,
Rome); Kansas (Tonganoxie State Lake); Kentucky
(Paducah); Maryland (Takoma Park); South Carolina
(Walhalla, Chappells, Florence); Texas (College Sta-
tion, Atlanta, Smith Co.); Washington, D.C. Riley
(1988:86) listed it from 6 parishes in Louisiana.
This is a wide and spotty distribution which needs
clarification by further collections. Many of the older
literature records for tristis possibly refer to this species.
Luginbill and Painter (1953: fig. 19) show tristis from
the eastern two thirds of the U.S.

FLORIDA DISTRIBUTION (fig. 474): The only
definite records are from Alachua Co. and Union Co.
Older Florida records of tristis by Blatchley (1929:70)
from Haulover and Ft. Barrancas need to be reexam-
ined in the light of the several species now in the tristis
complex. Young & Thames (1949:128-129) recorded
tristis as "not uncommon around Gainesville' (proba-
bly all apicata) and "in western parts of the state"
(probably true tristis).

BIOLOGY & ECOLOGY: Reinhard (1941:526-532)
published the only biology data on this species (at
College Station, Texas) and compared it with other
members of the tristis complex: egg production aver-

y6d 57 (20 n87) train the first 6 6 l7f Aprd 0 the Phy86phag6 b862 8 Chapin
(fig.6 66, 12686,269,2n,640,426,427,457.
459, 461,476479)

8 6677 7 Phy 8 Chapi 1932 203.
Stage -8g6622days6 (197625) Thead87676form Cinationh6688678. Sailor 1942:159.
in October, but spend the winter in the p71 "n' Phyltsphaga (C07766.7'6) b6266. Sa.demen
eireng in Marhthfolowin 6. 6 Thlifcycle 1951:250.
for I ,,n and apm-oi, .1 1- i s he south, is Us,
shecesofayUS~secis. odffen--orte-TYPE LOCALITY: "Sitaag. dc In Vegas. His
q- wete found between the,, secies, Because of 'noss... cub
this~andth ... affisn linnuairbngtheir identity, romy
S' DIAGNOSIS. This speies i lated ad mamil. to
-1vra Cub-n sporires m thic saligero C -ronhi,
Adult H- P101W T here rno host dam reported '

In C'd .. d,,d .. an muvod,,ed ormireatin cialfer, f'

ecause of past mf6.iN of the 2 speciesV species).
62068766876 7log 871686 668766 766 ndpn

t Immolate, Alth uttdi~ the .9 rdp.tlpup.a

-ldd an, be esprd 1. show iiusitcabke diffeciciaes.
Both thefrst and thid moor lav-,i deutifieA as 1-Ir,
_cr d.afibed by Boviag (194':33, flg. 33-36). No
1-lilty, data _c W resented for diesc 1-sve, and it is

could a (o eer s aiui aradi-dec alier it on,

SPECIMENS EXAMINED: uvs, 800. of which741
_er from Florid., of which 739 weic from A ch.u
C. P.-Iesntig 31 crnlterur. occuids A. ld %-ci
forfrm = urdi (Dade Co., 1934) is questionable.
since, thrue no other ecouhhim ...thirff i-iffe
"crunmpletedmi-Apc~pcini.2 Corifisi-nbenwe
roembers cifthe .-ci cmple army be spursiluhlef., r
11-1 yr-ani, nihidentifiionam, but all the aboee
record, h-1 b. eofied l ady.

SELECTED REFERENCES. -hihld, 1939 U 11.
'. 1 5305 2. 195049; RIs. 1-84 88, Is 6,95-78. mW 1.
si-I., 21.0 la 4 15 12-Nsnabld~ ~ im~B (m
_668,16 6 7 6 7

DESCRIPTION (fig. 476): Length: 7.8-10.3mm;
Width: 4.7-5.3mm. Shape: cylindrical, parallel. Color:
brownish testaceous, piceous to aeneous on pronotal
disk, elytral margin, and margin of elytral suture.
Vestiture: glabrous, shining. Antenna: 9-segmented;
male club shorter than stem. Clypeus: emarginate;
almost flat, median indentation deep and angulate;
margin abruptly reflexed. Tarsal Claws: slender, slightly
curved; tooth triangular, median, stouter than apical
portion. Male Posterior Tibial Spurs: lower movable,
two-thirds length of upper; longer spur twice length of
first tarsal segment. Abdomen: fig. 404 (venter). Female
Genitalia: fig. 269 (ventral), 272 (lateral). Male Geni-
talia: fig. 6 (caudal), 66 (ventral), 126 (dorsal), 186
(right lateral).

TAXONOMIC NOTES: This introduced Cuban spe-
cies has no synonyms. Saylor (1942:159) created the
genus Cnemarachis for most of the West Indian species
of Phyllophaga. Sanderson (1951) treated it as a
subgenus. Variation in the claspers was described and
illustrated by Woodruff (1961:15, fig. 25). It is related
to suturalis (Chevrolat), aeruginosa (Burmeister), micro-
soma Chapin, aeneotincta Chapin, and alquizara Chapin
(all Cuban species).
I examined the holotype male labelled: USNM
Type 43800, Cuba, Santiago de las Vegas, Havana, 14-
VI-1921, B. T. Barreto, and compared it with Florida
specimens. They are conspecific in all respects, including
the genitalia. The allotype female was described by
Woodruff (1961:16, fig. 11) with the following label
data: Miami, Florida, 28-IV-60, P. E. Briggs, blacklight
trap (FSCA). Because the female was not known to
Chapin when he originally described the species, and
because of the importance of the genitalia in establishing
the identity of both males and females, it was deemed
important to create the allotype as a point of reference.
Even though this was more than 30 years after the
original description, it complies with the original definition
of the term allotype (see Fernald, 1939).

U.S. DISTRIBUTION (fig. 479): This introduced
Cuban species is known in the U.S. only in Florida.
Agricultural inspectors (pers. com.) at the Arizona
border have intercepted it several times in vehicles
originating in Miami, Florida.

FLORIDA DISTRIBUTION (fig. 478): Originally
found only in Miami (Woodruff, 1959, 1960, 1961), it
has spread to 3 counties; Broward, Collier, and Dade
(see Appendix 3 for complete data).

BIOLOGY & ECOLOGY: Considerable work was
done on this Cuban species shortly after it was intro-
duced into Florida about 1959. It is now well estab-
lished, and in the 30 succeeding years it has spread to
both coasts. This species is a known pest of sugarcane
in Cuba, but it has not yet been found in the sugarcane
growing regions around Lake Okeechobee. It has been
the subject of a Master's thesis (Samol, 1968) and a 2
year USDA, ARS grant (Habeck & Wolfenbarger,
1968). The following resume' is taken from personal
observations, along with data from the above refer-
ences.
Egg production varies greatly, but averages about
12 per female. No specimens have been reared from
egg to adult, and no definite larval food has been
established. Presumably they feed on grass roots.
Adults are found during the day, within the top 3 inches
of soil; larvae in the upper 6 inches. Average head
capsule width for the 3 larval instars is as follows: 0.95,
1.56,2.47mm. Two peaks of adult activity occur (86%
April, May, June; 12% August, September, October),
although adults emerge every month of the year. The
greatest number taken on one night in a blacklight trap
was 58,400 on May 12, 1965. Adult evening flight
takes place primarily at about 25 minutes after sunset,
with less than 1 foot candle of light. Their return from

Adult Host Plants: Woodruff (1961: table 1) provided a list of host plants in Miami:

Family

Common Name

Bombacaceae
Juglandaceae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Malvaceae
Meliaceae
Moraceae
Phytolaccaceae
Polygonaceae
Sapindaceae
Sapindaceae
Sapotaceae
Sapotaceae
Sapotaceae
Ulmaceae

Pachira aquatica Aubl.
Carya illinoinensis (Wangenh.) K. Koch
Bauhinia sp.
Cassiafistula L.
Cassia marginata Roxb.
Castanospermum australe A. Cunn.
Delonix regia (Bojer) Raf.
Erythrina sp.
Gliricidia sepium Stend.
Gliricidia sp.
Tamarindus indica L.
Hibiscus rosa-sinensis L.
Swietenia mahagoni Jacq.*
Ficus roxburghii Wall.
Phytolacca rigida Small
Ruprechtia sp.
Euphoria longana Lam.
Melicoccus bijugatus Jacq.
Pouteria sapota (Jacq.) Moore & Steam
Pouteria viridis (Pittier) Cronquist
Chrysophyllum oliviforme L.*
Trema micrantha (L.) Blume*

(*Native Florida Plants)

the foliage to their hiding places occurs early in the
morning during a brief 15 minute period (6 to 6:15a.m.
on May 17-19). Duration of copulation averaged 21.6
minutes (17 to 26).
The giant Surinam toad, Bufo marinus Linnaeus,
fed on adults. The larvae of a luminescent click beetle,
Pyrophorus havaniensis Castelnau, were found feeding
on larvae. Adult click beetles of this species were
collected by the hundreds (it was previously rare) in the
same light traps with the Phyllophaga.
Within 2 years of its discovery (June, 1959) in
Miami, it had occupied an area of about 10 square
miles. It was not seen at the Subtropical Experiment
Station (near Homestead) until 1968. It was first found
at Ft. Lauderdale in 1977 and on the west coast at
Naples in 1986.
In addition to the list of positive hosts above, a list
of 36 negative hosts was provided (Woodruff, 1961:
table 2). The native Trema micrantha (L.) Blume
(=floridana) appears to be a preferred host for both the

v\ -W/ '

\ iI //

fig. 477. Phyllophaga bruneri: 10th abdominal segment (ventral),
third instar larva, note scattered setae not arranged in rows or
pallidium (see fig. 460) (13mm 1mm).

Genus and Species

shaving-brush tree
pecan
bauhinia
golden shower tree
senna
moreton-bay chestnut
Royal poinciana
coral-bean
madre de Cacao
madre
tamarind
Chinese hibiscus
mahogany

poke berry

longan
Spanish lime
sapote
green sapote
satinleaf
Florida trema