The flat bark beetles of Florida (Coleoptera: Silvanidae, Passandridae, Laemophloeidae)

THE FLAT BARK BEETLES OF FLORIDA
(COLEOPTERA:SILVANIDAE, PASSANDRIDAE, LAEMOPHLOEIDAE)

By

MICHAEL CHARLES THOMAS

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

UNIVERSITY OF FLORIDA

1985

ACKNOWLEDGMENTS

I would like to thank the following curators for their

cooperation and patience in making possible the many loans

that made this work possible: R.E. Woodruff (FSCA), T.J.

Spilman and G.H. Hevel (USNM), R.J.W. Aldridge (BMNH), N.

Berti (MNHN), D.H. Kavanaugh (CASC), A.F. Newton, Jr.

(MCZC), E.C. Becker (CNC), and S.A. Slipinski (PASC).

Individuals who supplied many valuable specimens were:

Gary Shook, Boise, ID, Albert Allen, Twin Falls, ID, Karl

Stephan, Red Oak, OK, R.H. Turnbow, Jr., Fort Rucker, AL,

and Tim King, Birmingham, AL. D.G.H. Halstead, Slough,

England, identified or confirmed my identifications for most

of the specimens of Silvaninae recorded here and supplied

specimens of many species. Ken Curry, University of Florida,

helped identify gut contents of many specimens. Miss Lori

Parker of Miami helped tend an ultraviolet light trap that

produced many important records.

The members of my committee, Drs. R.E. Woodruff, J.H.

Frank, D.H. Habeck, R.I. Sailer, and J. Reiskind, have

provided guidance, encouragement and valuable criticism, for

which I am deeply indebted. My chairman, R.E. Woodruff, has

been a reservoir of support and knowledge for many years.

Last, but certainly not least, I want to thank my wife,

Sheila. She has, more than anyone, made this work possible.

TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS...........................................ii

LIST OF FIGURES.................................... .... ......vi

ABSTRACT............... ................................... xi

INTRODUCTION..... ............. .............................1

MATERIALS AND METHODS....................................... 6

CHECKLIST OF FLAT BARK BEETLES FOR AMERICA NORTH OF MEXICO..

KEY TO THE GENERA OF CUCUJIDAE, SILVANIDAE, PASSANDRIDAE AND
LAEMOPHLOEIDAE KNOWN TO OCCUR IN AMERICA NORTH OF MEXICO..17

SILVANIDAE.................................................. 25

Uleiotinae .......................................... 28
Telephanani....... ............................. 29
Genus Cryptamorpha Erichson.......................... 30
Cryptamorpha desiardinsi (Gu6rin-Meneville)...........32
Genus Telephanus Wollaston ...........................34
Telephanus velox (Haldeman)...........................35
Silvaninae .......................................... 38
Genus Nausibius Redtenbacher.........................38
Nausibius clavicornis (Kugelann).....................40
Nausibius maior Zimmermann.............................42
Nausibius repandus LeConte...... .....................44
Nausibius sahlbergi Grouvelle........................46
Genus Eunausibius Grouvelle ..........................47
Eunausibius salutaris Parsons..........................49
Genus Oryzaephilus Ganglbauer..........................50
Genus Silvanus Latreille..............................54
Silvanus lewisi Reitter ..............................56
Silvanus muticus Sharp..... ...........................57
Silvanus planatus Germar...............................58
Silvanus proximus Grouvelle..... .....................60
Silvanus recticollis Reitter.........................62
Genus Cathartosilvanus Grouvelle.....................63
Cathartosilvanus imbellis (LeConte)..................64
Cathartosilvanus opaculus (LeConte)..................66
Genus Silvanoprus Reitter............................68
Silvanoprus scuticollis (Walker).....................70
Genus Monanus Sharp................................... 71
Monanus concinnulus (Walker).........................73
Genus Ahasverus Gozis.................................75
Ahasverus advena (Waltl) .............................77

Ahasverus longulus (Blatchley)........................80
Ahasverus rectus (LeConte) ...........................81
Genus Cathartus Reiche.................................82
Cathartus quadricollis (Gu6rin-M6neville).............84

PASSANDRIDAE................ ............................... 87

Genus Catogenus Westwood..............................89
Catogenus rufus (Fabricius) ..........................92
Genus Scalidia Erichson..... .........................93
Sclidia linearis (LeConte)..................... ....... 97

LAEMOPHLOEIDAE ............................................ 99

Genus Lathropus Erichson............................100
Lathropus pictus Schwarz... .........................101
Lathropus vernalis Casey...... .......................103
Genus Rhabdophloeus Sharp ...........................106
Rhabdophloeus horni (Casey) .........................108
Genus Cryptolestes Ganglbauer .......................109
Cryptolestes n.sp. 1..................................116
Crvptolestes n.sp. 2...................................119
Cryptolestes uncicornis (Reitter)...................121
Cryptolestes punctatus (LeConte)..................... 127
Genus Dysmerus Casey........ .........................129
Dysmerus basalis Casey ..............................132
Genus Leptophloeus Casey....... .....................133
Leptophloeus angustulus (LeConte)...................135
Genus Narthecius LeConte.............................138
Narthecius grandiceps (LeConte)......................139
Genus Laemophloeus Dejean............................141
Laemophloeus biguttatus (Say) ......................145
Laemophloeus fasciatus Melsheimer.....................147
Laemophloeus lecontei Grouvelle.....................148
Laemophloeus megacephalus Grouvelle.................150
Laemophloeus suturalis Reitter ......................153
Laemophloeus n.sp................................... 156
Genus Charaphloeus Casey ............................158
Charaphloeus adustus (LeConte)......................160
Charaphloeus bituberculatus (Reitter)...............162
Charaphloeus convexulus (LeConte)...................163
Genus Placonotus Macleay ............................164
Placonotus macrognathus Thomas......................167
Placonotus modestus (Say) ...........................169
Placonotus politissimus (Wollaston).................172
Placonotus zimmermanni (LeConte).....................174
Genus Parandrita LeConte & Horn.....................175
Parandrita permixtus (Grouvelle)....................176
Genus Phloeolaemus Casey ............................179
Phloeolaemus chamaeropis (Schwarz)..................181

SUMMARY...... .......... ..................................184

APPENDIX................................................ 204

REFERENCES CITED............................................ 218

BIOGRAPHICAL SKETCH....................................... 238

LIST OF FIGURES

Fig. 1. Cryptamorpha desiardinsi (Gu6rin-Mgneville), habitus

.......................................... ........32

Fig. 2. Telephanus velox (Haldeman), habitus..............36

Fig. 3. Nausibius repandus LeConte, habitus...............45

Fig. 4. Eunausibius salutaris Parsons, habitus............48

Fig. 5. Orvzaephilus acuminatus Halstead, habitus.........52

Fig. 6. Silvanus planatus Germar, habits .................59

Fig. 7. Cathartosilvanus imbellis (LeConte), habitus......65

Fig. 8. Silvanoprus scuticollis (Walker), habitus.........69

Fig. 9. Monanus concinnulus (Walker), habitus.............74

Fig. 10a. Ahasverus rectus (LeConte), habitus.............79

Fig. 10b. Ahasverus advena (Waltl), prothorax.............79

Fig. 10c. Ahasverus longulus (Blatchley), prothorax....... 79

Fig. 11. Cathartus quadricollis Gugrin-Mgneville, habitus...

.......................................... .......85

Fig. 12. Catogenus rufus (Fabricius), habitus.............91

Fig. 13. Scalidia linearis (LeConte), habitus.............96

Fig. 14. Lathropus pictus Schwarz, habitus...............102

Fig. 15. Rhabdophloeus horni (Casey), habitus............107

Fig. 16. Cryptolestes n.sp. 2, habitus...................118

Fig. 17. Cryptolestes uncicornis (Reitter), habitus......122

Fig. 18. Cryptolestes punctatus (LeConte), habitus.......126

Fig. 19. Dysmerus basalis Casey, habitus.................131

Fig. 20. Leptophloeus angustulus (LeConte), habitus......136

Fig. 21. Narthecius grandiceps (LeConte), habitus........140

Fig. 22. Laemophloeus biguttatus (Say), habitus..........146

Fig. 23. Laemophloeus lecontei Grouvelle, habitus........149

Fig. 24. Laemophloeus megacephalus Grouvelle, habitus....151

Fig. 25. Laemophloeus n. sp., habitus....................155

Fig. 26. Charaphloeus bituberculatus (Reitter), habitus..161

Fig. 27. Placonotus macrognathus Thomas, habitus.........168

Fig. 28. Placonotus modestus (Say), habitus..............170

Fig. 29. Parandrita permixtus (Grouvelle), habitus.......178

Fig. 30. Phloeolaemus chamaeropis (Schwarz), habitus.....180

Fig. 31. Orvzaephilus acuminatus Halstead, front tarsus.....

....................................... ........ 187

Fig. 32. Cathartosilvanus imbellis (LeConte), prosternum....

.................................... .............187

Fig. 33. Cucuius clavipes Fabricius, hind tarsus, male...187

Fig. 34. Uleiota dubius (Fabricius), aedeagus............187

Fig. 35. Uleiota dubius (Fabricius), pro-and mesosternum....

................................................ 187

Fig. 36. Catogenus rufus (Fabricius), pro-and mesosternum...

............. ........................ ...........187

Fig. 37. Cucujus clavipes Fabricius, pro-and mesosternum....

................................................189

Fig. 38. Charaphloeus convexulus (LeConte), pro-and

metasternum.......................................... 189

Fig. 39. Catogenus rufus (Fabricius), mouthparts, ventral

view ..................................... .......189

Fig. 40. Scalidia linearis (LeConte), hind tarsus....189

vii

Fig. 41. Catogenus rufus (Fabricius), hind tarsus....189

Fig. 42. Placonotus modestus (Say), pronotum.......... 189

Fig. 43. Placonotus modestus (Say), aedeagus, lateral view..

..................... ........................... 191

Fig. 44. Uleiota dubius truncatus Motschulsky, anterior

pronotal angle...................................191

Fig. 45. Uleiota dubius truncatus Motschulsky, mandible,

male ..................................... .......191

Fig. 46. Dendrophagus sp., hind tarsus....................191

Fig. 47. Cryptamorpha desiardinsi (Gu4rin-M&neville), head,

dorsal view..................................... 191

Fig. 48. Cryptamorpha desiardinsi (Gu6rin-M6neville), hind

tarsus............................................ 191

Fig. 49. Telephanus velox (Haldeman), maxillary palpus...191

Fig. 50. Telephanus velox (Haldeman), hind tarsus........193

Fig. 51. Cucuius clavipes Fabricius), head...............193

Fig. 52. Laemophloeus biguttatus (Say), intercoxal process..

sternum III..................................... 193

Fig. 53. Cryptolestes sp., intercoxal process sternum III...

............................. ................... 193

Fig. 54. Leptophloeus angustulus (LeConte), intercoxal

process sternum III..............................193

Fig. 55. Narthecius sp., head, dorsal view...........193

Fig. 56. Eunausibius salutaris Parsons, antennal cavity.....

.................................. ...........195

viii

Fig. 57. Cathartosilvanus imbellis (LeConte), femoral line,

sternum III..................................... 195

Fig. 58. Silvanus sp., femoral lines......................195

Fig. 59. Silvanus sp., anterior pronotal angle...........195

Fig. 60. Pensus gilae (Casey), anterior pronotal angle...195

Fig. 61. Silvanoprus scuticollis (Walker), femoral line.....

.......................... ....................... 195

Fig. 62. Cathartus quadricollis (Gu6rin-M6neville), antennal

club.................................... ..........195

Fig. 63. Abasverus advena (Waltl), antennal club.........195

Fig. 64. Ahasverus rectus (LeConte), antennal club.......195

Fig. 65. Oryzaephilus surinamensis (Linnaeus), temple....195

Fig. 66. Orvzaephilus mercator (Fauvel), temple..........195

Fig. 67. Cryptolestes punctatus (LeConte), aedeagus......197

Fig. 68. Cryptolestes uncicornis (Reitter), aedeagus.....197

Fig. 69. Cryptolestes ferrugineus (Stephens), genital

sclerite........................................ 199

Fig. 70. Cryptolestes ferrugineus (Stephens), sclerite of

bursa........................................... 199

Fig. 71. Cryptolestes pusillus (Schonherr), genital sclerite

................. ...... ................ .........199

Fig. 72. Cryptolestes pusillus (Schonherr), sclerite of

bursa .......................................... 199

Fig. 73. Cryptolestes klapperichi Lefkovitch, genital

sclerite.......................................... 199

Fig. 74. Cryptolestes klapperichi Lefkovitch, sclerite of

bursa ............................................ 199

Fig. 75. Cryptolestes n.sp. 1, aedeagus and internal sac....

................................................ 201

Fig. 76. Cryptolestes turcicus (Grouvelle), genital sclerite

.................................................. 201

Fig. 77. Cryptolestes turcicus (Grouvelle), sclerite of

bursa ............................................ 201

Fig. 78. Cryptolestes pusilloides (Steel & Howe), genital...

sclerite......................................... 201

Fig. 79. Cryptolestes pusilloides (Steel & Howe), sclerite

of bursa.......................................... 201

Fig. 80. Cryptolestes n.sp. 2, aedeagus and internal sac....

.................. ... ... ..... ......... ........203

Fig. 81. Cryptolestes n.sp. 2, sclerite of bursa..........203

Fig. 82. Laemophloeus fasciatus Melsheimer, left elytron....

............. .................... ............... 203

Fig. 83. Charaphloeus convexulus (LeConte), hind tarsus,

male ..................................... .......203

Fig. 84. Placonotus politissimus (Wollaston), femoral lines.

............................. ................. ... 203

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

THE FLAT BARK BEETLES OF FLORIDA
(COLEOPTERA:SILVANIDAE, PASSANDRIDAE, LAEMOPHLOEIDAE)

By

MICHAEL CHARLES THOMAS

August 1985

Chairman: R.E. Woodruff
Major Department: Entomology and Nematology

The Florida species of the cucujoid families

Silvanidae, Cucujidae, Passandridae, and Laemophloeidae are

reviewed. Keys and illustrations for all Florida genera and

species and diagnoses and detailed distribution records for

all non-economic species in the state are provided. A key to

the genera of the United States is also provided, as is an

current checklist. There are 14 genera and 33 species of

Silvanidae in the United States, 11 genera and 22 species of

which occur in Florida. Of the two U.S. genera and four

species of Cucujidae, none occur in Florida. Two genera and

two species of Passandridae occur in the U.S., including

Florida. The Laemophloeidae are represented by 14 genera and

53 species in the U.S., of which 11 genera and 31 species

are recorded from Florida. Two new species of Cryptolestes

and one of Laemophloeus are described. Newly recorded from

Florida are the following species: Nausibius sahlbergi

Grouvelle, Eunausibius salutaris Parsons, Monanus

concinnulus (Walker), Silvanus lewisi Reitter, S. proximus

Grouvelle, Laemophloeus suturalis Reitter, Charaphloeus

adustus (LeConte), Charaphloeus convexulus (LeConte),

Parandrita permixtus (Grouvelle), and Rhabdophloeus horni

(Casey). New combinations proposed are: Charaphloeus

bituberculatus (Reitter) (from Laemophloeus); Charaphloeus

flavosignatus (Schaeffer) (from Laemophloeus); Parandrita

permixtus (Grouvelle) (from Laemophloeus); Phloeolaemus

chamaeropis (Schwarz) (from Laemophloeus); Rhabdophloeus

horni (Casey) (from Cryptolestes); Rhabdophloeus disseptus

(Casey) (from Cryptolestes). Charaphloeus Casey and

Phloeolaemus Casey are elevated from subgenera of

Laemophloeus to full generic rank. Cryptolestes uncicornis

(Reitter) is revived from synonymy under C. punctatus

(LeConte). New synonyms proposed are: Cathartosilvanus

opaculus (LeConte) = C. trivialis (Grouvelle); Laemophloeus

megacephalus Grouvelle = L. floridanus Casey; Charaphloeus

convexulus (LeConte) = C. filiger Casey and C. sphaerops

Casey; Charaphloeus adustus (LeConte) = C. fraterculus

Casey; Cryptolestes uncicornis (Reitter) = C. quadratus

(Casey); Cryptolestes punctatus (LeConte) = C. adumbratus

Casey and C. extricatus Casey; Lathropus pictus Schwarz = L.

costatus Grouvelle. Lectotypes are designated for

Laemophloeus quadratus Casey, Cryptolestes adumbratus Casey,

Cryptolestes extricatus Casey, and Laemophloeus megacephalus

Grouvelle.

xiii

INTRODUCTION

The flat bark beetles are a diverse group of taxa

traditionally included in the family Cucujidae, but recently

considered to belong to three or four separate families.

They are ubiquitous under the bark of dead deciduous, and to

a lesser extent, evergreen trees and logs. Most species

appear to be fungivorous, feeding primarily on ascomycete

fungi. However, some species are found only in the galleries

of Scolytidae and may be an important, but little studied,

element in the natural control of bark beetles.

Like other members of the series Cucujiformia, the flat

bark beetles possess cryptonephritic Malpighian tubules,

making them well-adapted to relatively dry habitats and

preadapting them to life in stored products. Indeed, several

species are among the most destructive of stored products

pests and have been distributed throughout the world by

commerce.

Although the biology and taxonomy of the economically

important species are reasonably well-known, the same cannot

be said for other U.S. flat bark beetles. Many, if not most,

museum specimens are either not identified or misidentified,

and the only current identification keys deal strictly

with the stored products species. As Arnett (1962:776)

concluded, "The group is badly in need of study at all

levels." Because of this, I have included a checklist of

described United States species assigned to their proper

genus, and a key to the adults of genera known to occur in

the United States. Of the 32 genera and 92 species of these

families known to occur in the United States, 24 genera (75

percent) and 55 species (60 percent) occur in Florida, so

that the keys permit the identification of the majority of

the species in the United States.

The importance of knowing the taxonomy and biology of

apparently non-economic species is illustrated by two recent

examples: Orvzaephilus acuminatus Halstead was described in

1980 from specimens from India and Sri Lanka and from

coconut shells imported into Great Britain (Halstead 1980).

A year later, Jacob (1981) discussed the species' potential

as a stored products pest. Three years later, Thomas and

Woodruff (1984) recorded the importation of 0. acuminatus

into Florida from India. Cryptolestes klapperichi Lefkovitch

was described from Afghanistan in 1962 (Lefkovitch 1962a);

was recorded from Arabia in 1965 (Lefkovitch 1965b), and was

reported to be a stored products pest in Sri Lanka and

Malaysia (Green 1979). It is here reported from the New

World for the first time, based on specimens collected in

the wild in the U.S. Virgin Islands, and it is apparently

actively expanding its range, probably through the agency of

commerce.

3

The only published list of flat bark beetles from

Florida is that of Schwarz (1878), who recorded 13 species

of the families treated in the present work. Fifty-five

species are included in this study, including economic

species which may not be established in the state but which

have been, or are likely to be, recorded from Florida.

Economic species (especially those of Cryptolestes and

Oryzaephilus) which do not occur, or very rarely occur, in

the wild, are keyed and illustrated but are otherwise not

considered in detail. The taxonomic and biological

literature on such species is voluminous.

Although larval diagnoses are presented for each

family, this work is predominantly concerned with the

identification and taxonomy of adults. Statements about

specimens refer to adults unless otherwise stated to refer

to other stadia.

This study is based on more than 2,000 specimens

collected in Florida, deposited primarily in the Florida

State Collection of Arthropods (FSCA), Gainesville, and in

my personal collection (MCTC), plus more than a thousand

specimens from outside the state. Collection data for all

Florida specimens examined for this study are listed in the

Appendix. Types and other specimens were also examined from

the United States National Museum of Natural History,

Washington, D.C. (USNM), Field Museum of Natural History,

Chicago (FMNH), California Academy of Sciences, San

4

Francisco (CASC), British Museum (Natural History), London

(BMNH), Museum National d'Histoire Naturelle, Paris (MNHN),

Canadian National Collection, Ottawa (CNC), and Polish

Academy of Science, Warsaw (PASC).

With the increasing interest in habitat protection as a

method of environmental preservation, the distribution

records compiled during this study pinpoint those areas in

which isolated populations of West Indian species continue

to exist, and which face imminent threat of extinction from

the continued urbanization of southern Florida. Several of

these species may very well be useful as indicator species

of especially fragile habitats.

Woodruff (1973:x) noted that "Florida is an ideal state

for faunal studies because it is a peninsula and is

delimited on three sides. It is an important area from a

zoogeographic standpoint because of the proximity of the

West Indies and because of the high percentage of endemic

species."

Although this is intended primarily as a faunal study,

the taxonomy of these groups is such that numerous taxonomic

and nomenclatural changes have been necessary. To prevent

possible nomenclatural problems, the three new species

described in this work are not named. Recent revisions to

the International Code of Zoological Nomenclature (Anonymous

1985) restrict publication within the meaning of the Code to

works reproduced by ink on paper (Article 8c) and permit

5

authors to include a disclaimer that the work is issued for

the permanent scientific record (Article 8b). To prevent

problems from other nomenclatural changes made in this

dissertation, I here make that disclaimer.

MATERIALS AND METHODS

Flat bark beetles are more abundant in nature than

their representation in museum collections suggests, but

because of their usually minute size and retiring habits

they do not often come to the attention of the general

collector. Their small size also necessitates special

preparation in order to accurately observe their external

skeletal structure and genitalia.

Ultraviolet light-trapping, particularly in areas with

much dead wood, is especially productive and is the method

by which the majority of specimens for this study were

obtained. It is the most efficient method for gathering

basic data on the spatial and temporal distribution of the

adults. However, it reveals little information on their

habits and habitats.

In Florida, where my collecting experience has been

confined, most flat bark beetles can be found under the bark

of hardwood trees and logs, especially the various oaks,

Quercus spp. The stage of decay and physical location of the

log (e.g., whether it is located in the sun or shade,

whether it is contact with the ground, or is above the

ground) are important factors in determining the abdundance

of flat bark beetles and, probably, the composition of taxa

present. A few species (e.g., Phloeolaemus chamaeropis

(Schwarz)) are most abundant on freshly downed wood, but

most laemophloeines and silvanines can be found most

abundantly on logs that have been down long enough for the

bark to crack and ascomycete fungi to separate the thin

outer bark from the thick, corky layer of inner bark. It is

within these pockets of loose outer bark that many flat bark

beetles can be collected (as well as colydiids,

rhizophagids, nitidulids, mycetophagids, and corylophids).

(The fruiting bodies of ascomycete fungi are flat, black or

grey structures that look like splotches of tar and not at

all like the fleshy fungi with which most collectors are

familiar.) Flat bark beetles can also be found, although

less abundantly, in the space between the inner bark and the

sapwood on older, well-seasoned logs. Moist-rotten logs

rarely produce many specimens of flat bark beetles.

Most general entomology textbooks recommend a stout

knife or screwdriver for prying bark. I have found the best

tool for this task is a pry bar, available in many hardware

stores. This resembles a miniature crowbar but is fashioned

from a flat bar of tempered steel. Both ends are sharpened

and it can be used for chopping as well as prying. A small

pocket knife is useful for separating the thin outer bark

from the inner bark. An aspirator is essential, since these

beetles tend to scurry rapidly into crevices and under

attached bark when they are exposed, and are much too small

for rapid hand-collecting.

8

I have also collected specimens by beating dead

branches and by placing leaf debris in a modified Berlese

funnel. The latter method produces some silvanines, but few

other flat bark beetles. I have seen specimens of several

species that were collected in vane traps baited with

ethanol. This is a potentially productive collecting

procedure that should be investigated in greater detail.

Flat bark beetles can be killed in a standard killing

jar or in a vial of 70-75 percent isopropyl alcohol. Ethanol

probably should not be used since it seems to cause the soft

tissues of the body to harden and makes the clearing of

specimens difficult.

Most specimens can be glued to standard points, but

representative specimens should be mounted on microscope

slides. The following procedure has given me good results:

clear the specimen by boiling in a dilute (approximately 10

percent by weight) solution of potassium hydroxide (KOH) or

sodium hydroxide (NaOH). I use a small glass crucible and an

alcohol lamp, but a small beaker and hot plate would work as

well. Once the soft tissues have cleared (the specimen will

appear transparent but not colorless), rinse the specimen in

distilled water or a weak solution of acetic acid (vinegar

will do). Using fine pins or needles under a dissecting

microscope, dissect away the abdomen, and separate the

tergum from the sternum. The genitalia will come away with

the tergum. The mouthparts can be removed by placing the

point of the pin firmly against the mentum and pulling

anteriorly; this will usually separate all of the mouthparts

from the head capsule except for the mandibles, which can

then be pulled away with forceps or the tip of the pin. To

better observe the structure of the pro- and mesocoxal

cavities, the legs on one side of the body should also be

removed. If the body is heavily pigmented, it may be

bleached in a 3 percent solution of hydrogen peroxide.

Body, genitalia, and mouthparts should be mounted under

separate cover slips. I use Hoyer's solution as a mounting

medium so that specimens can be mounted directly from water or

alcohol; for greater permanence balsam may be used, but

requires time-consuming dehydration of specimens. Using 12mm

diameter cover slips, three mounts may be made on the same

slide, thus keeping the different parts of the specimen

together.

Larvae are often present in the same habitat as the

adults and should be collected and kept associated with the

adults. I use KAAD (1 part kerosene, 7 to 10 parts 95

percent ethanol, 2 parts glacial acetic acid, 1 part dioxane

(Peterson 1962:7)) larval fixative for killing larvae;

specimens are then stored in vials of 70 percent isopropyl

alcohol or are slide-mounted. Rearing larvae in Petri dishes

or snap-cap plastic vials with bits of fungus-covered bark

is relatively simple and would provide much needed

information on the immature stages of these beetles. The

10

larvae of most species, and even many Nearctic, genera are

unknown.

A representative of at least one species is illustrated

for each genus; it should be possible to identify many

specimens to genus by reference to the illustrations. The

illustrations were prepared using a binocular dissecting

microscope equipped with an eyepiece scale. Specimen

dimensions were measured at either 15x or 30x. The

measurements were transferred to graph paper, and a

preliminary pencil sketch made. After transferring the

pencil sketch, the drawing was inked and then shaded by one

of the following techniques: stippling, scratch board, or

coquille board.

CHECKLIST OF FLAT BARK BEETLES FOR AMERICA NORTH OF MEXICO

(Note: Species marked with an asterisk are present in or, in

the case of economic species, are likely to be found in

Florida. Where species are now assigned to a genus other

than the one in which they were originally placed the

original assignment is indicated in parenthesis after the

species name.)

SILVANIDAE
Uleiotinae

Uleiotini

Uleiota Latreille 1796:46
Brontes Fabricius 1801:97
Hyliota Reitter 1880:80
debilis LeConte 1856:76 (Brontes)
dubius Fabricius 1801:97 (Brontes)
truncatus Motschulsky 1845:92

Dendrophagus Schonherr 1809:50
cygnaei Mannerheim 1846:515
americanus Mannerheim 1853:207
glaber LeConte 1850:223
?germari Mannerheim 1843:306

Telephanini

Cryptamorpha Wollaston 1854:156
Pseudophanus LeConte 1860:84
desiardinsi Gu6rin-Mgneville 1829-44:196
(Psammoecus)
hubbardi Casey 1884:167 (Psammoecus)
musae Wollaston 1854:157
signatus LeConte 1860:85 (Pseudophanus)

Telephanus Erichson 1846:329
Heterodromia Haldeman 1848:127
lecontei Casey 1884:103
velox Haldeman 1848:127

Silvaninae

Nausibius Redtenbacher 1858:998
clavicornis Kugelann 1794:511 (Cucuius)
dentatus Marsham 1802:108 (Coricaria)
denticulatus Macleay 1825:46 (Sylvanus)
intermedius Smith 1851:16 (Silvanus)
latus Fairmaire 1850:53 (Silvanus)
parallelus Walker 1858:206 (Rhizophagus)
maior Zimmermann 1869:257
dentatus var. major, Crotch 1873:44
repandus LeConte 1866:70
sahlbergi Grouvelle 1896:206

Eunausibius Grouvelle 1912a:314
salutaris Parsons 1974:182

Orvzaephilus Ganglbauer 1899:584
mercator Fauvel 1889:132 (Silvanus)
gossypi Chittenden 1897:12 (Silvanus)
surinamensis Linnaeus 1758:357 (Dermestes)
bicornis Erichson 1846:337 (Silvanus)
frumentarius Fabricius 1775:62 (Anobium)
cursor Fabricius 1792:96 (Tenebrio)
sexdentatus Herbst 1783:31 (Ips)
sexdentatus Fabricius 1792:232 (Dermestes)
acuminatus Halstead 1980:309

Silvanus Latreille 1804:158
bidentatus Fabricius 1792:233 (Dermestes)
sulcatum Fabricius 1792:555 (Colydium)
affinis Reitter 1876:58
lewisi Reitter 1876:57
muticus Sharp 1899:560
nitidulus LeConte 1856:78
planatus Germar 1824:466
zimmermanni Guerin-Mgneville 1829-44:1
cognatus LeConte 1856:77
proximus Grouvelle 1904:183
amabilis Grouvelle 1914:140
recticollis Reitter 1876:61
?Leucohimatium breve Wollaston 1873-74::
reflexus Reitter 1880:25
vitulus Grouvelle 1882:294
pumilus Grouvelle 1912a:332
minimus Grouvelle 1912a:334
unidentatus Olivier 1790:9 (Ips)
planum Herbst 1797:285 (Colydium)
gratiosus Motschulsky 1863:501
siculus Stierlin 1864:147

98

170

Pensus Halstead 1973:77
gilae Casey 1884:72 (Silvanus)

Cathartosilvanus Grouvelle 1912a:340
imbellis LeConte 1856:77 (Silvanus)
communiss Grouvelle 1878:75 (Silvanus)
opaculus LeConte 1856:78 (Silvanus)
trivialis Grouvelle 1878:75 (Silvanus)

Silvanoprus Reitter 1911:45
angusticollis Reitter 1876:59 (Silvanus)
scuticollis Walker 1859:53 (Silvanus)
triangulus Reitter 1876:60 (Silvanus)

Monanus Sharp 1879:85
Emporius Ganglbauer 1899:578
concinnulus Walker 1858:207 (Monotoma)
signatus Frauenfeld 1867:438 (Silvanus)
fasciatus Wollaston 1873:169 (Cryptamorpha)
fascipennis Reitter 1876:129 (Cathartus)

Cathartus Reiche 1854:77
quadricollis Guerin-Meneville 1829-44:198
(Silvanus)
cassiae Reiche 1854:78
gemellatus Jacquelin du Val 1857:104
(Silvanus)

Ahasverus Gozis 1881:cxxvii
advena Waltl 1834:169 (Cryptophagus)
guerini Allibert 1847:12 (Silvanus)
musaeorum Ziegler 1844:270 (Latridius)
angustatus Lucas 1847:221 (Silvanus)
striatus Rouget 1876:ccvii (Crvptophagus)
longulus Blatchley 1910:564 (Cathartus)
parviceps Casey 1916:136 (Silvanus)
rectus LeConte 1856:78 (Silvanus)

CUCUJIDAE

Cucujus Fabricius 1775:204
clavipes Fabricius 1781:257
var. subnitens Schaeffer 1931:175
puniceus Mannerheim 1843:303

Pediacus Shuckard 1839:185
depressus Herbst 1797:286 (Cucuius)
subglaber LeConte 1856:73
fuscus Erichson 1846:313
dermestoides Schonherr 1809:53 (Cucuius)
plans LeConte 1850:223
subcarinatus Mannerheim 1852:363

PASSANDRIDAE

Catogenus Westwood 1835:221
Anisoceras Hope 1840:131
Anisoceras Gemminger and Harold 1868:869 (not Hope 1840)
rufus Fabricius 1798:123 (Cucuius)
puncticollis Newman 1839:399
parvus Casey 1916:116
puncticeps Casey 1916:116
monilicornis Casey 1916:115

Scalidia Erichson 1846:305
Taphroscelidia Crotch 1873:44
Syssitos Sharp 1899:541
linearis LeConte 1863:70 (Catogenus)

LAEMOPHLOEIDAE

Lathropus Erichson 1846:327
vernalis Casey 1884:95
pictus Schwarz 1878:358
costatus Grouvelle 1902:763
pubescens Casey 1884:96
striatus Casey 1916:119
robustulus Casey 1916:120

Rhabdophloeus Sharp 1899:531
horni Casey 1884:89 (Laemophloeus)
disseptus Casey 1916:133 (Cryptolestes)

Cryptolestes Ganglbauer 1899:612
Leptus Thomson 1863:92
Fractophloeus Kessel 1921:28
capensis Waltl 1834:169
clavicollis Wollaston 1854:479 (Laemophloeus)
elongatulus Lucas 1849:479 (Laemophloeus)
vermiculatus Wollaston 1854:164 (Laemophloeus)
rotundicollis Casey 1884:89 (Laemophloeus)
ferrugineus Stephens 1831:223 (Cucuius)
testaceus Paykull 1800:168 (not
Fabricius 1787) (Cucuius)
monilicornis Stephens 1831:223 (Cucuius)
concolor Smith 1851:6 (Laemophloeus)
obsoletus Smith 1851:7 (Laemophloeus)
carinulatus Wollaston 1877:44 (Laemophloeus)
emgei Reitter 1887:286 (Laemophloeus)
alluaudi Grouvelle 1906:120 (Laemophloeus)
pubescens Casey 1884:93 (Laemophloeus)
punctatus LeConte 1856:75 (Laemophloeus)
adumbratus Casey 1916:134
extricatus Casey 1884:92 (Laemophloeus)
geminatus LeConte 1856:75 (Laemophloeus)

pusilloides Steel and Howe 1952:86 (Laemophloeus)
pusillus Schonherr 1817a:55
minutus Olivier 1791:243 (not Fourcroy
1785) (Cucuius)
testaceus Stephens 1831:224 (Cocuius)
crassicornis Waltl 1839:225
(Laemophloeus)
longicornis Mannerheim 1843:303
(Laemophloeus)
brevis Fairmaire 1850:56 (Laemophloeus)
parallels Smith 1851:7 (Laemophloeus)
pauper Sharp 1899:530 (Laemophloeus)
puberulus LeConte 1856:75 (Laemophloeus)
turcicus Grouvelle 1876:xxxii (Laemophloeus)
truncatus Casey 1884:93 (Laemophloeus)
weisei Reitter 1880:78 (Laemophloeus)
schwarzi Casey 1884:83 (Laemophloeus)
uncicornis Reitter 1874:45 (Microbrontes)
recticollis Reitter 1876:52 (Laemophloeus)
denticornis Casey 1884:94 (Laemophloeus)
iteratus Sharp 1899:528 (Laemophloeus)
addendus Sharp 1899:529 (Laemophloeus)
quadratus Casey 1884:90 (Laemophloeus)
n.sp. 1
n.sp. 2

Leptophloeus Casey 1916:135
Truncatophloeus Kessel 1921:28
angustulus LeConte 1866:379 (Laemophloeus)

Dysmerus Casey 1884:97
Brontophloeus Kessel 1921:28
basalis Casey 1884:97

Narthecius LeConte 1861:95
Paraphloeus Sharp 1899:509
grandiceps LeConte 1863:70
breviceps Casey 1890:323
monticola Fall 1907:222
oregonensis Hatch 1961:200
simulator Casey 1890:322
striaticeps Fall 1907:222

Laemophloeus Dejean 1836:340
biguttatus Say 1827:267 (Cucuius)
bisignatus GuBrin-Meneville 1829-44:205
californicus Casey 1916:122
fasciatus Melsheimer 1846:113
fervidus Casey 1916:121
megacephalus Grouvelle 1876:425
floridanus Casey 1884:85
lecontei Grouvelle 1876:496

shastanus Casey 1916:123
suturalis Reitter 1876:50
terminalis Casey 1884:83
n.sp. 1

Charaphloeus Casey 1916:127
adustus LeConte 1856:74 (Laemophloeus)
fraterculus Casey 1916:124
bituberculatus Reitter 1876:316 (Laemophloeus)
convexulus LeConte 1879:2 (Laemophloeus)
filiger Casey 1916:125
sphaerops Casey 1916:126
dimidiatus Schaeffer 1910:215 (Laemophloeus)
flavosignatus Schaeffer 1910:214 (Laemophloeus)

Metaxyphloeus Thomas 1984a:67
texanus Schaeffer 1904:201 (Rhinomalus)

Placonotus Macleay 1871:168
Silvanophloeus Sharp 1899:537
arizonensis Thomas 1984c:7
macrognathus Thomas 1984c:12
modestus Say 1827:268 (Cucuius)
singularis Smith 1851:7 (Laemophloeus)
liguidus Casey 1916:130 (Silvanophloeus)
gundlachi Grouvelle 1874:499 (Laemophloeus)
nitens LeConte 1856:75 (Laemophloeus)
bullatus LeConte 1856:75 (Laemophloeus)
politissimus Wollaston 1867:67 (Laemophloeus)
mirus Grouvelle 1905:142 (Laemophloeus
(Silvanophloeus))
victus Kessel 1926:69 (Laemophloeus
(Silvanophloeus))
commixtus Grouvelle 1912b:304 (Laemophloeus)
zimmermanni LeConte 1856:75 (Laemophloeus)
apertus Casey 1916:128 (Silvanophloeus)
illustris Casey 1916:129 (Silvanophloeus)
sobrinus Casey 1916:129 (Silvanophloeus)

Parandrita LeConte and Horn 1880:133
cephalotes LeConte 1856:76 (Laemophloeus)
permixtus Grouvelle 1912b:303 (Laemophloeus)

Phloeolaemus Casey 1916:127
chamaeropis Schwarz 1878:359 (Laemophloeus)

Notolaemus Lefkovitch 1959a:102
macrocephalus Schaeffer 1910:214 (Laemophloeus)

Deinophloeus Sharp 1899:540
impressifrons Schaeffer 1910:213 (Laemophloeus)

KEY TO THE GENERA OF CUCUJIDAE, SILVANIDAE, PASSANDRIDAE AND
LAEMOPHLOEIDAE KNOWN TO OCCUR IN AMERICA NORTH OF MEXICO

1. Tarsomere I longer than II (fig. 31), tarsal

formula 5-5-5 in both sexes; aedeagus with

parameres on dorsal aspect of median lobe;

procoxal cavities closed posteriorly, usually

broadly (fig. 32); suture between mesosternum and

mesepisternum incomplete or obsolete (Silvanidae:

Silvaninae) .....................................23

1'. Tarsomere I usually shorter than II (fig. 33),

tarsal formula 5-5-5 or 4-4-4 in both sexes, or 5-

5-4 in males only; aedeagus with parameres on

ventral aspect of median lobe (fig. 34); procoxal

cavities usually open posteriorly (fig. 35, 36,

37, 38); suture between mesosternum and

mesepisternum usually complete...................2

2(1'). Genae produced anteriorly as rounded plates

covering maxillae (fig. 39); elytra, but not

pronotum, with longitudinal grooves; tarsal

formula 5-5-5 in both sexes; mesocoxal cavities

closed laterally by juncture of meso- and

metasterna (fig. 36) (Passandridae)..............3

18

2'. Genae not produced anteriorly, maxillae free; if

elytra with longitudinal grooves then pronotum

also with grooves or carinae; tarsal formula often

5-5-5 in both sexes or 5-5-4 in males; mesocoxal

cavities usually closed laterally by mesepisternum

or mesepimeron or both (fig. 35, 37, 38).........4

3(2). Body subcylindrical (fig. 13); tarsomere I as long

as II (fig. 40)..................Scalidia Erichson

3'. Body dorsoventrally compressed (fig. 12);

tarsomere I distinctly shorter than II (fig. 41)..

................................. Catogenus Westwood

4(2'). Pronotum with longitudinal lines paralleling

lateral margins (fig. 42); parameres fused to

basal piece and reduced to tooth-like processes on

ventral aspect of aedeagus (fig. 43)

(Laemophloeidae)................................10

4'. Pronotum without longitudinal lines paralleling

lateral margins; parameres freely articulated to

basal piece and not reduced (fig. 34)............5

5(4'). Antennal scape elongate (fig. 2); tarsi not

heteromerous in males, tarsomeres lobed in some

genera (Silvanidae:Uleiotinae)....................6

5'. Antennal scape not elongate; tarsi heteromerous in

males, tarsomeres not lobed (Cucujidae)..........9

6(5). Procoxal cavities open posteriorly (fig. 35);

tarsomeres not lobed (Uleiotini) .................7

6'. Procoxal cavities closed posteriorly; tarsomeres

distinctly lobed (Telephanini)....................8

7(6). Tarsal formula 4-4-4; pronotum irregularly toothed

laterally and with anterior angles strongly

produced (fig. 44); males with curved mandibular

processes (fig. 45) ..............Uleiota Latreille

7'. Tarsal formula 5-5-5, basal segment short but

distinct (fig. 46); pronotum simple laterally,

anterior angles not produced; males without

mandibular processes........Dendrophagus Schonherr

8(6'). Head dorsally with longitudinal impressed lines

(fig. 47); apical maxillary palpomeres not

securiform; tarsomere III bilobed (fig. 48).......

............................Cryptamorpha Wollaston

8'. Head dorsally without impressed lines; apical

maxillary palpomeres securiform (fig. 49);

tarsomere III simply lobed (fig. 50)..............

.............................. Telephanus Erichson

9(5'). Head produced laterally behind eyes (fig. 51);

antennae not distinctly clubbed; color bright

red .............................. Cucujus Fabricius

9'. Head not produced laterally behind eyes; antennae

distinctly clubbed; color not bright red..........

................................. Pediacus Shuckard

10(4). Sublateral line of pronotum carinulate, obviously

raised above the surface of the pronotum;

transverse epistomal groove absent, although

epistome may be depressed........................17

10'. Sublateral line of pronotum a groove, sometimes

with a weakly expressed associated ridge; or, disc

of pronotum depressed but without obvious groove

or carinulate line; transverse epistomal groove

sometimes present ...............................11

11(10'). Intercoxal process of sternum III acuminate

apically (fig. 52)...............................12

11'. Intercoxal process of sternum III not acuminate

apically......................................... 14

12(11). Tarsomere I shorter than penultimate tarsomere;

labrum shallowly emarginate apically; elytra

carinate laterally.............Laemophloeus Dejean

12'. Tarsomere I at least subequal to penultimate

tarsomere; labrum semicircular, not emarginate

anteriorly; elytra usually not carinate

laterally........................................ 13

13(12'). Head rostrate.................Metaxyphloeus Thomas

13'. Head not rostrate...............Charaphloeus Casey

14(11'). Head with distinct transverse groove separating

frons from epistome (fig. 27, 28).................

................................. Placonotus Macleay

21

14'. Head without distinct transverse groove separating

frons from epistome, although region of head

corresponding to clypeus may be depressed below

level of frons...................................15

15(14'). Anterior coxal cavities wide open posteriorly;

mandibles of males in some species expanded

laterally (fig. 30).............Phloeolaemus Casey

15'. Anterior coxal cavities nearly closed posteriorly;

mandibles of males not expanded laterally.......16

16(15'). Elytral epipleura well defined, complete to apical

angle; head moderate in size; basal angles of

epistome not foveate; major males with clypeal

horns ...........................Deinophloeus Sharp

16'. Elytral epipleura poorly defined, becoming

obsolete at midpoint of elytra; head large, almost

as large as pronotum (fig. 29); basal angles of

epistome foveate; males without clypeal horns.....

.................................Parandrita LeConte

17(10). Lateral margins of pronotum slightly to strongly

undulating (fig. 14, 15); disc sometimes with

paired anterior and posterior depressions (fig.

14, 15), these often faint.......................18

17'. Lateral margins of pronotum evenly curved, or, at

most, simply sinuate; disc without paired anterior

and posterior depressions........................19

22

18(17). Head with lateral lines; lateral margins of

pronotum with irregular undulations; pronotum and

elytra explanate laterally, elytra carinate

laterally and with longitudinal discal costae;

anterior coxal cavities open posteriorly;

intercoxal process of sternum III narrowly rounded

anteriorly (fig. 15)...........Rhabdophloeus Sharp

18'. Head without lateral lines; lateral margins of

pronotum undulating at most; elytra without

elytral cells, not carinate; intercoxal process of

sternum III broadly rounded anteriorly (fig. 14)..

.................................Lathropus Erichson

19(17'). Anterior coxal cavities open posteriorly; elytra

with only third cell developed....................

............................. Notolaemus Lefkovitch

19'. Anterior coxal cavities closed or nearly closed;

elytra with all three cells developed...........20

20(19'). Intercoxal process of sternum III broadly rounded

anteriorly (fig. 53)........Cryptolestes Ganglbauer

20'. Intercoxal process of sternum III narrowly rounded

anteriorly (fig. 54) ............................21

21(20'). Epistome acuminate anteriorly (fig. 55)...........

.................... ............. Narthecius LeConte

21'. Epistome truncate or emarginate anteriorly......22

23

22(21'). Pedicel attached laterally to scape, which is

large and irregularly produced in males (fig. 20);

pronotum quadrate...................Dysmerus Casey

22'. Pedicel attached axially to scape, which is normal

in both sexes; pronotum quadrate to elongate......

.................................Leptophloeus Casey

23(1). At least tarsomere III lobed ....................29

23'. No tarsomeres lobed.............................. 24

24(23'). Lateral margins of pronotum simple; hind femora in

males simple. .................................. 27

24'. Lateral margins of pronotum dentate or undulating;

hind femora in males toothed ....................25

25(24'). Lateral margins of pronotum undulating (fig. 3, 4)

.............. ........................... ........26

25'. Lateral margins of pronotum dentate (fig. 5)......

....................... Oryzaephilus Ganglbauer

26(25). Head ventrally with antennal cavity (fig. 56).....

.............................Eunausibius Grouvelle

26'. Head ventrally without antennal cavities..........

.................... ........ Nausibius Redtenbacher

27(24). Anterior angles of pronotum not produced

anterolaterally (fig. 7); femoral lines open (fig.

57) ....................Catbartosilvanus Grouvelle

27'. Anterior angles of pronotum produced

anterolaterally; femoral lines closed (fig. 58)...

..... .......... ................... ............... 28

24

28(27'). Anterior angles of pronotum acute, usually

produced more laterally than anteriorly (fig. 59).

................................ Silvanus Latreille

28'. Anterior angles of pronotum obtuse, produced more

anteriorly than laterally (fig. 60)...............

....................................... Pensus Halstead

29(23). Anterior angles of pronotum acute (fig. 8);

femoral lines closed (fig. 61)....................

..............................Silvanoprus Grouvelle

29'. Anterior angles of pronotum lobed at most; femoral

lines closed or open ............................30

30(29'). Pronotal margins simple; femoral lines open;

elytra immaculate............................... 31

30'. Pronotal margins denticulate (fig. 9); femoral

lines closed; elytra maculate........Monanus Sharp

31(30). Body elongate, parallel-sided (fig. 10);

antennomere XI at its broadest narrower than X

(fig. 62); intercoxal process of sternum III

pointed anteriorly................Cathartus Reiche

31'. Body ovate (fig. 11); antennomere XI at its

broadest equal in width to X (fig. 63, 64);

intercoxal process of sternum III rounded

anteriorly .........................Ahasverus Gozis

SILVANIDAE

Diagnosis. The combination of the following character

states will distinguish adults of this family from the

others treated in this work: anterior coxal cavities closed

in most genera (open in Uleiota and Dendrophagus, which are

not present in Florida); aedeagus with articulated

parameres; tarsal formula 5-5-5 in both sexes (4-4-4 in some

species of Uleiota); antennae either clubbed or filiform,

not moniliform; mandibles with dorsal mycangium (a more or

less circular cavity on the dorsal surface of the mandible

near the base the function of which, in at least one

silvanid, is the transport of fungal spores (Crowson and

Ellis 1969)); maxillae exposed, not concealed by genal

processes; head and pronotum without sublateral carinae or

grooves; elytra without elytral cells but with scutellary

striole.

Larval diagnosis. Elongate, parallel-sided, somewhat

flattened; mandible with acute retinaculum, and well-

developed mola; mala acute apically, cardines bipartite;

five or six pairs of stemmata; antennae either with three

well-developed antennomeres (Uleiotinae) or with antennomere

III reduced and fused to antennomere II (Silvaninae);

frontal suture somewhat lyriform; frontoclypeal suture

absent; dorsal surface of thoracic and abdominal segments

26

without asperities; spiracles annular; legs well-developed,

two tarsungular setae, close together in Silvaninae, widely

separated in Uleiotinae; long, whip-like urogomphi present

in some Uleiotinae, absent in all Silvaninae; abdominal

segment X located terminally, produced as pseudopod.

Taxonomic notes. Like the other families treated in

this work, this family has until recently been usually

treated as a subfamily of the Cucujidae. However, Crowson

(1955:104) removed the former subfamilies Silvaninae and

Psammoecinae to the Silvanidae. Crowson (1967:211) later

expressed doubt, stating "...Passandrinae and Silvaninae

could well be subfamilies of Cucujidae..."; however, Sen

Gupta and Crowson (1966:63; 1969:586-590) and Crowson

(1973:62) continued to treat the Silvanidae, including the

subfamily Uleiotinae, as a full family. Thomas (1984b:47-54)

reviewed the phylogeny and suprageneric classification of

the Silvanidae and divided it into two subfamilies,

Uleiotinae and Silvaninae, with the former composed of two

tribes, Uleiotini and Telephanini. In a similar study, Pal

et al. (1985:213-217), retained Cryptamorphinae and

Psammoecinae as subfamilies, in addition to Silvaninae and

Uleiotinae, which they, like Thomas (1984b), considered to

exhibit the most ancestral character states among the

Silvanidae.

Hetschko (1930) listed 34 genera and 391 species

worldwide that would now be assigned to Silvanidae. Genera

27

and species described, and synonymies proposed since then

(primarily by Nevermann, Halstead, Pal, and Sen Gupta),

bring the world total to about 47 genera and approximately

470 species.

Biology. The biology and immature stages of the great

majority of species and most genera are unknown, but the

following generalizations can be made (the known biology of

each genus and species occurring in Florida is reviewed

below): Uleiotinae (Uleiotini) are found primarily under

bark, where both adults and larvae probably feed on

ascomycete and other fungi (Crowson and Ellis 1969; personal

observations); Uleiotinae (Telephanini) are found primarily

on plants, especially withered, pendant leaves, and in plant

debris, where they also probably feed predominantly on

fungi; Silvaninae are found under bark (e.g., Silvanus,

Cathartosilvanus) or in leaf litter or soil (e.g.,

Silvanoprus, Monanus, some Ahasverus), where they also seem

to feed on fungi. Some silvanines have been reported to be

at least facultatively predaceous. Several species belonging

to Orvzaephilus, Nausibius, Cathartus, and Ahasverus are

important pests of stored grains, grain products, nuts, and

spices. Their taxonomy and biology are the best known of the

Silvanidae.

Distribution. The family is worldwide in distribution,

but is most abundant at both the generic and species level

in the tropics. There are 32 species here recorded from the

28

United States and 22 from Florida, including several stored

products species. Although they may not be established, they

may be introduced repeatedly.

Uleiotinae

Diagnosis. Members of the Uleiotinae may be

distinguished from members of the Silvaninae by the

following combination of character states: aedeagus inverted

with parameres located on the ventral aspect of the median

lobe; tarsomeres in some genera distinctly lobed ventrally;

antennae elongate and filiform, without a distinct club.

Taxonomic notes. The traditional definition of the

Uleiotinae includes only Uleiota, Dendrophagus, and, by

extension, Brontopriscus. This is the arrangement followed

by Hetschko (1930) and Pal et al. (1984). Thomas (1984b)

grouped both the Uleiotini (composed of Uleiota,

Dendrophagus, Brontopriscus, and Aplatamus) and the

Telephanini (composed of Telephanus, Cryptamorpha, Platamus,

Indophanus, and Psammoecus) in the Uleiotinae. Until

recently (Crowson 1973), the Uleiotinae had been included in

the Cucujidae rather than the Silvanidae.

Biology. Reviewed above.

Distribution. Worldwide but more abundant in the

tropics. For example, Telephanus is represented by two

species in the United States and about 100 in the

Neotropics. Aplatamus and Platamus are exclusively

Neotropical; Cryptamorpha is confined to the Old World

tropics, except for one tropicopolitan species; Psammoecus

is predominantly Old World tropical, but with species in

Europe, Japan and northern Asia; Telephanus is almost

exclusively Neotropical, with two U.S. species, and several

described from the Old World tropics; Indophanus is confined

to India; and Brontopriscus is confined to New Zealand. Four

genera and seven species are known to occur in the U.S.; two

of these occur in Florida. Only the tribe Telephanini is

represented in Florida.

Telephanini

Diagnosis. From members of Uleiotini (which is not

known to be represented in Florida), those of Telephanini

can be distinguished by the following combination of

character states: anterior coxal cavities closed; tarsomere

I longer than II; antennal scape either very elongate (fig.

2) and maxillary palpi securiform (fig. 49) or head with

longitudinal grooves (fig. 47); pronotum laterally with

spines or stout setae.

Taxonomic notes. Is reviewed in this section under the

subfamily.

Biology. Members of this tribe are found on plants or

in leaf litter (see section on biology under subfamily).

Distribution. Worldwide but most abundant in the

tropics (see this section under subfamily).

Genus Cryptamorpha Wollaston

Cryptamorpha Wollaston 1854:156

Pseudophanus LeConte 1860:84; syn. by Waterhouse 1877:122

Parabrontes Redtenbacher 1867:40; syn. by Waterhouse 1877:122

Type species. Of Cryptamorpha, Psammoecus desiardinsi

Guerin-Meneville [by synonymy of Cryptamorpha musae

Wollaston]; of Pseudophanus, P. signatus LeConte [by

monotypy]; of Parabrontes, P. silvanoides Redtenbacher [by

monotypy].

Diagnosis. Cryptamorpha most resembles Telephanus in

the New World, but can be distinguished easily by the

longitudinal grooves on the frons, shorter antennal scape,

non-securiform maxillary palpi, and strongly bilobed

tarsomere III.

Taxonomic notes. The generic synonymy was first

proposed by Waterhouse (1877:122) and has been accepted by

most subsequent authors. Hetschko (1930:88) listed 24

species in this genus; Pal and Sen Gupta (1979:78-80)

described two new species from India and Bhutan. The genus

has never been revised, nor are there any comprehensive keys

to the species. Most of the original descriptions are brief

and without illustrations, thus specific identifications

require, in most cases, comparisons with types.

Biology. The little information published on the

biology of this genus (almost entirely on Cryptamorpha

31

desiardinsi) suggests that their biology is similar to

members of Telephanus.

Distribution. Of the species listed by Hetschko

(1930:88-90), only desiardinsi and redtenbacheri (Reitter)

occur in the New World. The former is tropicopolitan; the

latter may be generically distinct (Thomas 1984b).

Cryptamorpha desiardinsi (Guerin-Meneville)

(fig. 1, 47)

Psammoecus desiardinsi Guerin-Meneville 1829-44:196

Cryptamorpha musae Wollaston 1854:157; syn. by Waterhouse

1877:122

Pseudophanus signatus LeConte 1860:85; syn. by Waterhouse

1877:122

Psammoecus hubbardi Casey 1884:167; syn. by Casey 1890:498

Diagnosis. Length 3.3-4.4mm. Cryptamorpha desiardinsi

can only be confused with Telephanus velox in the Florida

fauna; the generic diagnoses will distinguish the two

species. According to the key in Grouvelle (1917:45), it

can be distinguished from other species of Cryptamorpha by

possession of prosternal process not inflexed behind the

coxae and elytra less than three times longer than wide.

Taxonomic notes. Because of its wide distribution, this

species has been described several times in several genera.

Waterhouse (1877:122) proposed most of the above synonymies.

Casey (1890:498) synonymised his own species. Adults appear

to be distinct and easily distinguished from adults of the

Fig. 1. Cryptamorpha desiardinsi (Gugrin-Meneville),
habitus.Length 3.3mm 4.4mm.

other species of the genus by the characters given in the

diagnosis above.

Biology. Blatchley (1928:67) reported collecting a

specimen at Dunedin, Florida in December ". .. by beating a

pile of dead leaves of cabbage palmetto. ." Loding

(1945:79) reported it "At light and in mud on river bank."

Lepesme (1939:62) reported it was imported into France on

bananas and pineapples (". presque exclusivement sur les

bananes et les ananas avec lesquels il est imports. .").

He also reported that adults feed on plant debris, but that

the larvae are predaceous ("L'adulte parait se nourrir de

debris d'origine veg6tal. La larve, au contraire, est

carnassiere ."). Bowler et al. (1977:452), however,

reported that larvae were abundant in Hawaii beneath leaf

sheaths of sugarcane infected with sugarcane smut (Ustilago

scitaminea Syd.), upon which they feed.

Distribution. Hetschko (1930:88) listed it as

"Kosmopolit". Blackwelder (1945:423) recorded it from St.

Vincent and Grenada. LeConte's Pseudophanus signatus was

described from Puget Sound, Ore. As Casey (1884:105) and

Batch (1961:206) pointed out, this was most likely based on

an introduction; it has not been reported since from the

Pacific Northwest. Leonard (1928:382) reported it from New

York City; this record also is almost certainly based on an

introduced specimen. Loding (1945:79) recorded it from

Alabama.

Specimens examined. Twenty-eight, all from Florida,

representing 11 collection records (for complete data see

Appendix).

Genus Telephanus Erichson

Telephanus Erichson 1846:329

Heterodromia Haldeman 1848:127; syn. by Melsheimer 1853:45

Diagnosis. 3.6mm 4.3mm. The laterally spinose

prothorax (fig. 2), elongate antennal scape, securiform

maxillary palpi (fig. 49), inverted aedeagus, and

approximate mesocoxae are diagnostic for members of this

genus.

Taxonomic notes. The date for Haldeman's paper has

usually been cited as 1846; however, the title page to the

volume indicates it was actually printed in 1848. This is

one of the largest silvanid genera, with Hetschko (1930)

listing 64 species. Nevermann (1931, 1932, 1937) added 47

species from Central America and the Antilles, and Thomas

(1984b) added a new species from Jamaica. There are only two

species known from the United States. Although the Central

American and Antillean species are fairly well-known because

of Nevermann's revisions, the South American fauna is in

need of revision; South American specimens of Telephanus are

virtually unidentifiable. Specimens should be handled

carefully, since the easily-broken prothoracic spines are

important in identification.

35

Biology. Nevermann (1931) presented a detailed review

of the biology of the Neotropical members of this genus.

Like most members of this tribe, these beetles appear to be

associated with withered, pendant leaves or litter rather

than loose bark. I collected numerous specimens of

Telephanus haitianus Nevermann by beating Lobelia assurgens

in the Massif de la Selle mountains in southeastern Haiti.

Distribution. This is a predominantly New World genus,

with 105 of the 114 described species occurring in the New

World, where members of the genus range from southern Canada

south to Brazil and throughout the Antilles. Species have

also been described from Mauritius, New Guinea, Madagascar,

Borneo, and Hawaii.

Telephanus velox (Haldeman)

(fig. 2, 49)

Heterodromia velox Haldeman 1848:127

Telephanus velox, Melsheimer 1853:45

Diagnosis. Length 3.6-4.3mm. The generic characters

given in the diagnosis above distinguish it from all other

Florida silvanids.

Taxonomic notes. This is such a distinct species that

there has been no confusion over its identity. Erichson

(1846:329-330) described the genus in a footnote and

proposed the new species name atricapillus. Since Erichson

did not describe the species, this name has been treated as

Fig. 2. Telephanus velox (Haldeman), habitus. Length
3.6mm 4.3mm.

a nomen nudum in most subsequent literature (although it was

treated as a valid name by Smith (1851:14)).

Biology. Casey (1884:103) noted that it is ". .. very

common under rubbish of various sorts ." Smith

(1910:263) reported that in New Jersey it occurs "Throughout

the State, under stones and old leaves; rarely under bark;

may be sifted out from fall to late spring." Blatchley

(1910:569) wrote that in Indiana it "Occurs more often

beneath stones, chunks and dead leaves than beneath bark.

When exposed it usually remains quiescent with antennae

folded against sides, but if touched it runs with great

swiftness, whence its specific name." Kirk (1969, 1970)

recorded it in South Carolina from "woodstrash," broomsedge,

in hollow trees, at lights, and on soil. The Florida

specimens I have seen were collected in ultraviolet light

traps, in pitfall trap, and from soybean plants.

Distribution. LeConte (1856:77) reported it from the "

S. Middle, Southern and Western States ." It has also

been recorded from New Jersey (Smith 1910:263), North

Carolina (Brimley 1938), New York (Leonard 1928), Alabama

(Loding 1945), Indiana (Blatchley 1910), South Carolina

(Kirk 1969, 1970), and Florida (Thomas 1979). I have seen

specimens from Arizona, Delaware, Florida, Georgia,

Illinois, Indiana, Iowa, Maryland, Massachusetts, Michigan,

New Hampshire, New York, North Carolina, Ohio, Oklahoma,

38

Ontario, Tennessee, Virginia, and Wisconsin. In Florida it

has been collected only in the northern third of the state.

Specimens examined. More than 80, of which 18 were from

Florida, representing four collection records (for complete

data see Appendix).

Silvaninae

Diagnosis. Members of this subfamily can be

distinguished by the following combination of character

states: tarsal formula 5-5-5 in both sexes; tarsomere I

longer than II; mandibular mycangium present, but reduced in

size compared to that found in Uleiotinae; antennae clubbed;

anterior coxal cavities closed; aedeagus not inverted,

parameres located on dorsal aspect of median lobe.

Taxonomic notes. Although until recently placed as a

subfamily of the Cucujidae, this group of genera is distinct

and easily characterized.

Biology. The biology of the members of this subfamily

were reviewed in this section under the family discussion.

Distribution. Worldwide, but most abundant and diverse

in the tropics.

Genus Nausibius Redtenbacher

Nausibius Redtenbacher 1858:998

Type species. Cucuius clavicornis Kugelann [by synonymy

of Corticaria dentata Marsham, type by monotypy].

Diagnosis. Most similar in Florida to Orvzaephilus and

Eunausibius. Members of Nausibius can be separated from

39

those of Oryzaephilus by having tarsomere III simple

(incrassate in Oryzaephilus) and pronotum laterally undulate

instead of toothed. The antennal club in Nausibius also

appears to be composed of four antennomeres, and pronotum

usually with a crescentic depression at base.

Taxonomic notes. According to Halstead (1980:344) this

genus, as presently composed, is polyphyletic, with

repandus, sahlbergi, and similar species generically

distinct from the group of species related to major and

clavicornis.

Biology. The biology of most species is unknown.

Nausibius clavicornis is a minor pest of stored products,

especially raw or yellow crystal sugar (Breese and Wise

1959); N. major is found under bark of oaks (Quercus spp.),

and N. repandus has been collected on pine (Pinus spp.) in

association with bark beetles (Scolytidae).

Distribution. Except for the cosmopolitan N.

clavicornis, members of this genus are restricted to the New

World.

Key to the adults of Florida species of Nausibius

1. Body depressed, slender, parallel-sided; pronotal

margins slightly undulating; pronotum basally

without distinct crescentic depression ...........2

40

1'. Body more convex, more robust, not as parallel-

sided; pronotal margins distinctly undulating;

pronotum basally with distinct crescentic

depression. ...................................... 3

2(1). Coarsely, densely punctate and pubescent dorsally,

punctures deep; color piceous...repandus (LeConte)

2'. Dorsal surface appearing almost impunctate,

punctures very shallow and ill-defined, dorsal

surface glabrous; color castaneous................

................................. .sahlbergi Grouvelle

3(1'). Pronotum with a distinctly less densely punctate

area medially (cosmopolitan, stored products pest)

.............................clavicornis (Kugelann)

3'. Pronotum without a distinctly less densely

punctate area medially............maior Zimmermann

Nausibius clavicornis (Kugelann)

Cucujus clavicornis Kugelann 1794:511

Nausibius clavicornis, Fauvel 1889:132

Corticaria dentata Marsham 1802:108; syn. by Fauvel 1889:132

Sylvanus denticulatus Macleay 1825:46; syn. by Halstead

1980:345

Silvanus intermedius Smith 1851:16; syn. by Gemminger and

Harold 1868:880

Silvanus latus Fairmaire 1850:53; syn. by Fauvel 1884:73

Rhizophagus parallelus Walker 1858:206; Hetschko 1930:73

41

Diagnosis. Length 3.2mm 4.1mm. Adults of this

species can be distinguished from the very similar major by

the sparser punctation of the pronotal disc.

Taxonomic notes. Halstead (1980:349) noted that many

North American literature records for this species actually

refer to major, with which it has long been confused.

Biology. Halstead (1980:349) wrote that it is ". .

undoubtedly subcorticolous in the field .", and

Ganglbauer (1899:581) reported that it is found in old nests

of bees in South America. Breese and Wise (1959) reviewed

the literature records and reported in detail on its biology

in the laboratory. They list the following commodities in

which clavicornis has been found: rice, dried apples,

ginger, logwood (Haematoxylon), and cassia husks. It is most

common in sugar, especially those kinds of unrefined sugars

with a high proportion of molasses, but Breese and Wise

(1959:257) concluded: "It is unlikely that N. clavicornis

could multiply rapidly in raw or unrefined sugar, and any

deterioration in stored sugar directly attributable to it

would probably be small in comparison with that caused by

other factors." Wolcott (1950:298) reported it from Puerto

Rico and noted it ". has repeatedly been found since in

brown sugar."

Distribution. Halstead (1980:347) wrote that "Although

indigenous to the Neotropical region, this species has been

carried to all parts of the world by commerce and has become

42

established in warmer regions." He reported specimens from

Madeira, Zaire, Uganda, Zimbabwe, Reunion, Sri Lanka, Burma,

China, Japan, Philippines, Hawaii, Australia, Canada,

U.S.A., Mexico, Guatemala, Belize, Nicaragua, Panama,

Brazil, Cuba, Jamaica, Puerto Rico, Dominica, St. Vincent,

and Trinidad. In the U.S., he reported it from California,

Texas, New York, Pennsylvania, Maryland, North Carolina,

South Carolina, and Florida.

Specimens examined. Only three, and none from Florida.

Nausibius maior Zimmermann

Nausibius maior Zimmermann 1869:257

Nausibius dentatus var. major, Crotch 1873:44

Nausibius clavicornis auctorum (in part), not Kugelann 1794:511

Nausibius major, Halstead 1980:349

Diagnosis. Length 3.9mm 5.1mm. Adults have denser

punctation of the thoracic disc than those of clavicornis.

Halstead (1980:349) also stated that antennomere VIII is

less transverse and pronotal teeth less prominent in maior

than in clavicornis.

Taxonomic notes. Zimmermann (1869:257) noted the close

resemblance of his species to clavicornis, but wrote that

maior is ". somewhat larger, of black color, with

reddish-brown antennae and legs, and perceptibly more

coarsely punctured." Crotch (1873:44) treated major as a

variety of dentatus. Casey (1884:75) wrote, after examining

specimens of major, ". that it is simply N. dentatus

43

[=clavicornis] without any difference at all, as far as I

can discover." After being treated as a variety or synonym

of clavicornis for nearly a century, this name was revived

by Halstead (1980:349), who stated that ". I am

convinced that the magnitude of difference shown by external

characters warrants specific status for N. major."

Biology. Zimmermann (1869:257) said ". found only

under the bark of old oaks." Halstead (1980:350) reported

one specimen ". collected on 'Scarlet oak assoc

carpenter worm attack' and two were caught at light."

Although most of the specimens I have seen were without data

or were collected in ultraviolet light traps, I collected

several specimens from an oak (Quercus sp.) with a slime

flux.

Distribution. Zimmermann (1869:257) described it from

"Carolina". Halstead (1980:350) reported specimens from

Arizona, Arkansas, Colorado, District of Columbia, Florida,

Georgia, Illinois, Kentucky, Louisiana, Maryland, Michigan,

New Jersey, New York, North Carolina, Tennessee, Texas, and

Virginia. I have seen additional specimens from the states

of Ohio and Oklahoma. There is also a specimen in the FSCA

with the following data: MEXICO, Nuevo Leon, 4 mi. S.

Monterrey nr. Siesta Motel, 12-VII-1963 R.H. Arnett, Jr. and

E.R. VanTassell. This is the first record from Mexico and

the first from outside the U.S.A.

44

Specimens examined. More than 30, of which three were

from Florida, representing two collection records (for

complete data see Appendix).

Nausibius repandus LeConte

(fig. 3)

Nausibius repandus LeConte 1866:70

Diagnosis. Length 2.4mm 2.8mm. Adults of this

species are most similar to those of sahlbergi in the

Florida fauna, but differ in their much more coarsely

punctured and distinctly pubescent dorsal surface. Adults of

both repandus and sahlbergi differ from those of major and

clavicornis by their smaller size, more parallel-sided and

flattened body form, and lack of a basal crescentic

depression on the pronotum.

Taxonomic notes. This is a distinctive species with no

specific synonyms. As noted above, Halstead (1980:344 and in

litt.) stated that it and several other species, including

sahlbergi, are not congeneric with clavicornis.

Biology. Blatchley (1918:423) reported collecting a

specimen ". at Dunedin while sweeping huckleberry and

other low shrubs ." and (1928:67) ". beaten from

the tops of recently felled pine." On two occasions, I have

collected adults of this species from beneath outer bark

scales of small, cut pines and pine branches infested with

scolytids and one first instar silvanid larva from within

the bark beetle galleries themselves.

Fig. 3. Nausibius repandus LeConte, habitus. Length 2.4mm -
2.8mm.

46

Distribution. LeConte (1866:379) described it from

Washington, D.C. and Schwarz (1878:445) recorded it from

Florida. I have not been able to find any additional

published distribution records. I have seen specimens from

Florida and Alabama.

Specimens examined. Twenty, of which 18 were from

Florida, representing four collection records (for complete

data see Appendix).

Nausibius sahlbergi Grouvelle

Nausibius sahlbergi Grouvelle 1896:206

Diagnosis. Length 3.0mm 3.3mm. Similar to repandus

in general body form, but dorsally glabrous and with very

shallow, ill-defined punctures that give the impression that

the surface is almost impunctate. Additionally, all

specimens I have seen of sahlbergi have been castaneous in

color, those of repandus piceous.

Taxonomic notes. This species has only been mentioned

in the literature once since its description, and that was

Halstead's (1980:344) opinion that this and related species

are not congeneric with clavicornis and maior.

Biology. Nothing has been published on its biology.

Most specimens I have seen were collected in ultraviolet

light traps.

Distribution. It was described from Brazil; I have seen

specimens from Panama and Florida.

47

Specimens examined. More than 50, of which two were

from Florida, with the following label data: 2, "Dade Co.

Fla. v. 1949"/ "Cathartus quadricollis det. H.F. Strohecker"

(FSCA).

Genus Eunausibius Grouvelle

Eunausibius Grouvelle 1912a:314

Type species. Grouvelle (1912a:314) described this

genus for Nausibius tenebrionoides Grouvelle and N.

elongatus Grouvelle, without designating a type species.

Since Halstead is currently revising this group of genera, I

will leave it to him to designate a type species.

Diagnosis. The species are most similar to those of

Nausibius related to repandus, but they possess ventral

antennal grooves on the head (fig. 56).

Taxonomic notes. Except for the assignment of three

additional species by Schwarz and Barber (1921:192) and

Parsons (1974:181-184), there has been little taxonomic

activity in this genus. The species appear to be relatively

uncommon in collections.

Biology. Wheeler (1921:88-91) reported on the biology

of E. wheeleri Schwarz and Barber in Guyana. It inhabits the

hollow petioles of the ant tree, Tachigalia sp., where all

stages feed on the tissue of the inner walls of the petiole

and on honeydew, which the beetles actively solicit, from

coccids that also inhabit the petioles.

Fig. 4. Eunausibius salutaris Parsons, habitus. Length
3.3mm 4.1mm.

Distribution. Except for the single specimen from

Florida, the members of this genus appear to be restricted

to South America.

Eunausibius salutaris Parsons

(fig. 4, 56)

Eunausibius salutaris Parsons 1974:182

Diagnosis. Length 3.3mm 4.1mm. The well-marked

antennal grooves (fig. 56) and general habitus (fig. 4)

easily distinguish this species in the Florida fauna.

Taxonomic notes. Parsons (1974:181) noted that it and

E. lophius Parsons key to E. elongatus (Grouvelle) in

Grouvelle's (1912a:314) key, but both can be distinguished

from that species by their depressed and laterally carinate

elytra (cf. convex and not carinate). Eunausibius salutaris

can be immediately distinguished from lophius by its much

smaller eyes and by the antennal insertions, which are

situated one-third of the length of an eye in front of the

eyes in lophius and the length of an eye in salutaris.

Halstead (in litt.) questioned the generic assignment of

salutaris.

Biology. Nothing has been published on the biology of

this species. I have examined specimens from Venezuela that

were collected in association with Pseudococcidae.

Distribution. It was described from Brazil (Parsons

1974:184) and has not been reported in the literature since.

It is here recorded from Florida and Venezuela for the first

time.

Specimens examined. Four, of which one was from

Florida, with the following data: "FLORIDA: Broward Co. Ft.

Lauderdale V-1983 Coll. by F.W. Howard" (FSCA), and three

were from Venezuela: "Est. Exb. Yaritagua Edo Yaracuey 1967

Osorio Coll."/"asociados con Pseudococcidae en ca8a de

azdcar" (FSCA).

Genus Oryzaephilus Ganglbauer

(fig. 5)

Silvanus (Oryzaephilus) Ganglbauer 1899:584

Oryzaephilus, Reitter 1911:45

Type species. Silvanus surinamensis (Linnaeus) [by

subsequent designation of Halstead 1980].

Diagnosis. Of Florida silvanids, only members of

Oryzaephilus possess six distinct teeth on the lateral

margin of the prothorax (fig. 5). Additionally, members of

this genus can be distinguished by the combination of the

following character states (Halstead 1980:276-277): body

elongate, more or less parallel-sided; tarsomere III

incrassate; head without ventral grooves; antennae with 11

antennomeres, antenna with antennomeres IX-XI forming a

distinct club, antennomere XI not obviously acuminate;

temples present; femoral lines not produced caudally.

Taxonomic notes. With Halstead's (1980) revision, this

has become one of the taxonomically best known of flat bark

beetle genera. One of the prime reasons for this is their

economic importance as stored-products pests, especially O.

surinamensis, the saw-toothed grain beetle, and 0. mercator,

the merchant grain beetle. Described as a subgenus of

Silvanus by Ganglbauer in 1899, it was raised to generic

rank by Reitter (1911:45). Hetschko (1930:67-70) listed

eight species; Halstead (1980) included 13 species. Major

references to the taxonomy and morphology of members of the

genus include Grouvelle (1912a), Guillebeau (1890),

Honomichl (1978), Howe (1953), Joshi (1976), Pajni and Bedi

(1975), Slow (1958), and Spilman (1960).

Biology. Their natural habitat appears to be under bark

(Halstead 1980:274), although Thomas (1984b:53) suggested

that their occurrence under bark may be incidental to other

factors. References that deal with the biology of members of

this genus include: Aitken (1966), Arbogast (1976), Ashman

and Higgs (1968), Back and Cotton (1926), Barnes and

Kaloostian (1940), Blackman (1966), Corbett et al. (1937),

Howe (1956), Jacob (1981), Loschiavo (1976), Pierce et al.

(1981), Saxena and Kaul (1976), Sinha (1965), Surtees

(1963), White and Sinha (1981), Wojcik (1969).

0. surinamensis and 0. mercator are common and serious

pests of stored products, with surinamensis being more

abundant on cereals and cereal products, dried fruit, copra,

nuts, and carob, whereas mercator is more common on oil nuts

and their products (Halstead 1980:290, 309). Two other

Fig. 5. Oryzaephilus acuminatus Halstead, habitus. Length
3.3mm 3.7mm.

53

species, 0. gibbosus Aitken and 0. acuminatus Halstead, have

been intercepted in Great Britain on coconut shells (Aitken

1965; Halstead 1980). The latter species was collected

recently in large numbers in Florida in a shipment of neem

(Azadirachta indica A. Juas) seed imported from India

(Thomas and Woodruff 1984) and may be established.

Distribution. Except for the widely distributed stored

products species, the members of this genus are restricted

to the Old World, especially to the warmer areas. Three

species have been collected in Florida. Although both

surinamensis and mercator are virtually worldwide in

distribution, only the former is capable of surviving

temperate zone winters but cannot complete development at

temperatures below 20 degrees Celsius (Halstead 1980:290,

309). See Aitken (1975) for detailed distribution records.

Key to the adults of Florida species of Oryzaephilus

1. Temple at least one-half length of eye (fig. 65),

not sharply angulate behind.......................

............................surinamensis (Linnaeus)

1'. Temple less than one-third length of eye (fig.

66), sharply angulate behind......................2

2(10). Anterior pronotal angle moderately developed in

large males, body less elongate (3.6-4.1:1);

pronotum of large males with lateral ridges not

strongly elevated .................mercator Fauvel

54

2'. Anterior pronotal angles strongly developed in

large males (fig. 5); body more elongate (3.9-

4.4:1); pronotum of large males with lateral

ridges strongly produced.......acuminatus Halstead

Genus Silvanus Latreille

Silvanus Latreille 1804:158

Leptus Duftschmidt 1823:156 [replacement name for Silvanus]

TypeR species. Is unidentata Olivier [by monotypy].

Diagnosis. The species of Silvanus can be distinguished

in Florida by the following combination of character states:

lateral margins of pronotum simple; anterior angles produced

anterolaterally (fig. 6,59); tarsomere III simple.

Taxonomic notes. The taxonomic history of Silvanus is

similar to that of Laemophloeus in the Laemophloeidae. As

traditionally constituted, it was a highly polyphyletic

assemblage of silvanids sharing little more than a vague

external similarity. Grouvelle (1912a) recognized this and

proposed a number of subgenera, none of which was accepted

until recently. Hetschko (1930:59-63) included 55 species in

Silvanus. Halstead (1972) adopted three of the four

subgenera proposed by Grouvelle (1912a) and elevated them to

generic rank, in addition to proposing three new genera. As

defined by Halstead (1973), Silvanus contains 17 species and

is a more natural, easily characterized taxon than it was

previously.

55

Biology. These beetles are generally found under the

bark of logs and dead trees, where they are at least partly

fungivorous (Halstead 1973:42). Eight species of Silvanus

have been recorded from stored products imported into Great

Britain, but none is a serious pest; Halstead (1973:42)

noted that their presence ". may often be due to cross-

infestation from dunnage and timber."

Distribution. This genus is represented throughout the

world except for South America.

Key to adults of Florida species of Silvanus

(Modified from Halstead 1973)

1. Pronotum more or less parallel-sided, at least in

apical half; anterior angles obtusely lobed, not

at all acute; temples rounded .....................

............................... recticollis Reitter

1'. Pronotum not obviously parallel-sided, more or

less strongly convergent to base; anterior angles

more or less angulate, not lobed; temples

toothed.......................................... 2

2(1'). Anterior angles of pronotum less acute, not

strongly produced laterally (fig. 6).............3

2'. Anterior angles of pronotum more acute, strongly

produced laterally (fig. 59)..................... 4

56

3(2). Head and pronotum glossy, punctation of frontal

triangle obviously different from sides of head;

temple extending laterally beyond eye (fig. 6)

(common, widespread species under bark)............

...................................... planatus Germar

3'. Head and pronotum dull; punctation of frontal

triangle not obviously different from sides of

head; temple not extending laterally beyond eye

(single southern Florida record)..................

................................ proximus Grouvelle

4(2'). Eyes very large, separated dorsally by 1.5 to 1.7

times their length; pronotum without well-defined

lateral foveae ......................lewisi Reitter

4'. Eyes not as large, separated dorsally by 1.9 to

2.1 times their length; pronotum with well-defined

lateral foveae .... ...................muticus Sharp

Silvanus lewisi Reitter

Silvanus lewisi Reitter 1878:57

Diagnosis. Length 2.1mm 2.3mm. The very large eyes

distinguish adults of this species from those of all other

Florida species of Silvanus.

Taxonomic notes. This is a distinctive species with no

recorded synonyms.

Biology. Often imported into Great Britain on stored

products or dunnage from the Oriental region and Africa,

this species has been recorded on desiccated coconut,

57

tapioca flour, rice, pulses, bark residues, gum arabic,

groundnut kernels, and red beans, as well as under bark,

on freshly fallen coconut palms, and at light (Halstead

1973).

Distribution. Halstead (1973) listed the distribution

of S. lewisi as India, Sri Lanka, China, Vietnam, Taiwan,

Japan, West Malaysia, Singapore, Philippines, Java, Borneo,

New Guinea, Solomon Islands, Australia, Congo, and Ghana. It

has not been recorded previously from the New World; the

specimens listed below may represent separate introductions.

It is possible that this species has become established in

southern Florida.

Specimens examined. Four, of which two were from

Florida with the following data: 1, "Dade Co., Fla. 19 "

(FSCA); 1, "FLA., Broward Co., Davie 24-II-1983 L.

Daigle"/"on book in office" (FSCA).

Silvanus muticus Sharp

(fig. 59)

Silvanus muticus Sharp 1899:560

Diagnosis. Length 2.3mm 3.0mm. In Florida, only

adults of S. lewisi could be confused with those of this

species; both have acute, well-developed anterior pronotal

angles. However, in S. muticus the eyes are smaller and the

longitudinal pronotal depressions are distinct.

Taxonomic notes. This is a distinctive species with no

recorded synonyms. However, Halstead (1973:70) noted that it

has often been confused with the similar S. bidentatus,

which also occurs in North America, but apparently much less

frequently that does S. muticus.

Biology. Halstead (1973:70) recorded it from under bark

of pine, maple, oak, chestnut, and juniper, and at light.

Distribution. The type locality is Guatemala and

Halstead (1973:70) recorded it from Canada (Montreal), and

in the United States from Alabama, Arkansas, California,

Florida, Georgia, Illinois, Iowa, Kansas, Louisiana,

Maryland, Massachusetts, Michigan, New Hampshire, New

Jersey, New York, North Carolina, Ohio, Pennsylvania, South

Carolina, Tennessee, Texas, Virginia, West Virginia, and

Wisconsin.

Specimens examined. More than 120, of which 40 were

from Florida, representing 17 collection records (for

complete data see Appendix).

Silvanus planatus Germar

(fig. 6)

Silvanus planatus Germar 1824:466

Silvanus zimmermanni Guerin-M6neville 1829-44:198

Silvanus cognatus LeConte 1856:77

Diagnosis. Length 2.3mm 2.8mm. From adults of other

Florida species, those of S. planatus can be distinguished

by the following combination of character states: Anterior

Fig. 6. Silvanus planatus Germar, habitus. Length 2.3mm -
2.8mm.

pronotal angles obtuse (fig. 6); head and pronotum glossy;

longitudinal pronotal depressions absent (fig. 6).

Taxonomic notes. Adults of this species are very

similar to the European S. unidentatus (Olivier), which is

established in North America, and S. nitidulus LeConte.

Neither species is known to occur in Florida. From the

former, adults of planatus can be distinguished by their

glossy dorsal surface and longer temple, and from the latter

by the presence of basal pronotal angles and generally

larger body size.

Biology. Under bark of various hardwoods (Halstead

1973:65).

Distribution. Halstead (1973:65) recorded it from

Ontario and Quebec in Canada and, in the United States, from

Alabama, Arkansas, Florida, Georgia, Illinois, Indiana,

Iowa, Kansas, Louisiana, Maryland, Michigan, New Hampshire,

New Jersey, New York, North Carolina, Ohio, Oklahoma,

Pennsylvania, South Carolina, Tennessee, Texas, and

Virginia.

Specimens examined. More than 120, of which 77 were

from Florida, representing 17 collection records (for

complete data see Appendix).

Silvanus proximus Grouvelle

Silvanus proximus Grouvelle 1904:183

Silvanus amabilis Grouvelle 1914:140; syn. by Halstead

1973:61

61

Diagnosis. Length 1.8mm 2.3mm. The following

combination of character states distinguishes adults of this

species from those of other Florida species: Head and

pronotum dull; anterior pronotal angles gradually developed,

broad and blunt; elytral apex often toothed.

Taxonomic notes. Adults of proximus most resemble those

of planatus in the Florida fauna, but the dorsal surface is

dull rather than glossy as in the latter species.

Biology. "Occurs occasionally in association with

stored products imported into Britain from Africa and one

specimen was found on brazil-nut residues in a ship carrying

nuts from Belon, Brazil. It has been sifted from oil-palm

fruits, Elaeis caught at light and found in humus

and forest-floor litter." (Halstead 1973:63)

Distribution. Widespread in Africa and introduced into

the Neotropics. Halstead (1973:63) recorded it from Guinea,

Sierra Leone, Ivory Coast, Ghana, Cameroun, Gabon, Zaire,

South West Africa, Tanzania, Mexico, Venezuela, Brazil, St.

Thomas, Martinique, and Grenada. It is here recorded from

Florida for the first time.

Specimens examined. Six, of which five were from

Florida, with the following data: "FLORIDA: Broward County

1-1984 in a building Collected by Jacques" (FSCA).

Silvanus recticollis Reitter

Silvanus recticollis Reitter 1876:61

?Leucohimatium breve Wollaston 1873:170; see Halstead

1973:55

Silvanus reflexus Reitter 1880:25; syn. by Halstead 1973:56

Silvanus vitulus Grouvelle 1882:294; syn. by Halstead 1973:56

Silvanus (Microsilvanus) pumilus Grouvelle 1912a:332; syn.

by Halstead 1973:56

Silvanus (Microsilvanus) minimus Grouvelle 1912a:334; syn.

by Halstead 1973:56

Diagnosis. Length 1.7mm 2.2mm. Adults of this

species can be distinguished from those of other Florida

species of this genus by the following combination of

character states: Sides of pronotum more or less parallel to

apex; anterior angles obtuse and directed more anteriorly

than laterally; eyes small; dorsal surface dull.

Taxonomic notes. A widespread, variable species, it was

described several times by Reitter and Grouvelle, whose

species were synonymized by Halstead (1973:56). Grouvelle

(1912a:332) described Microsilvanus as a subgenus of

Silvanus for this species (and its synonyms). Halstead

(1973) did not employ subgenera in his revision of Silvanus

and related genera, and the status of Microsilvanus has not

been resolved. Halstead (1973:55) wrote that Leucohimatium

breve Wollaston, described from Japan, is "...almost

63

certainly the same species as recticollis...", but was

unable to locate Wollaston's type.

Biology. Nothing is known about the biology of this

species except that it is attracted to light.

Distribution. Widespread in the Old World tropics, it

has been recorded from India, Thailand, Vietnam, Laos,

Sulawesi, Japan, Ryukyu Islands, Congo, Zaire, Zimbabwe,

(Halstead 1973:57). Thomas (1979:357) reported it for the

first time from the New World, based on a Florida specimen.

There was some doubt whether the specimen was adventitious,

but a specimen subsequently collected in Georgia (see below)

suggests that this species is established in the

southeastern United States.

Specimens examined. Three, two of which were from the

United States with data as follows: 1, "FLA., Indian River

Co., Fellsmere 4-IX-1975 Coll. M.C. Thomas blacklight trap"

(MCTC); 1, "Clarke Co., GA. Whitehall Forest VI-21-1979 R.

Turnbow blk. light" (RHTC).

Genus Cathartosilvanus Grouvelle

Silvanus (Cathartosilvanus) Grouvelle 1912a:340

Cathartosilvanus, Halstead 1973:81

Type species. Silvanus opaculus LeConte [by synonymy of

Silvanus trivialis Grouvelle, type by monotypy].

Diagnosis. Members of this genus most resemble

those of Silvanus and can be distinguished by the following

combination of character states: different form of anterior

pronotal angles, which are located posterior to anterior

margin of pronotum, and femoral lines open.

Taxonomic notes. Grouvelle (1912a:340) described this

as a subgenus for trivialis; Halstead (1973:81) raised it to

generic rank.

Biology. Primarily subcortical in habitat.

Distribution. Restricted to the New World, where three

species are primarily Neotropical (one is restricted to the

Galapagos Islands; one penetrates the Nearctic in the

southwestern United States and in Florida), and one is

widespread in the Nearctic.

Cathartosilvanus imbellis (LeConte)

(fig. 7, 32, 57)

Silvanus imbellis LeConte 1856:77

Cathartosilvanus imbellis, Halstead 1973:82

?Silvanus communis Grouvelle 1878:75; see Halstead 1973:82

Diagnosis. Length 2.5-2.9mm. The following combination

of character states distinguish individuals of this species

from those of opaculus: length, 2.4mm 2.8mm.; pronotum

elongate.

Taxonomic notes. The synonymy of communis (Halstead

1973:82) was based on the original description and

illustration (Grouvelle 1878:75); Halstead was unable to

locate the type, which was from Brazil. Grouvelle

(1912a:373) suggested that Gugrin's (1829-44:198)

description of Silvanus (=Cathartus) quadricollis applied

Fig. 7. Cathartosilvanus imbellis (LeConte), habitus. Length
2.3mm 2.8mm.

best to this species, but this has not been followed by

subsequent authors.

Biology. Found almost exclusively under bark,

in Florida especially of Quercus spp.

Distribution. Halstead (1973:84) reported it from

"CANADA: Ontario and Quebec. U.S.A.: Oregon, Nebraska, Iowa,

Illinois, Kansas, Texas, Louisiana, Michigan, Indiana, Ohio,

New York, Massachusetts, Pennsylvania, W. Virginia,

Maryland, New Jersey, N. Carolina, Alabama, Georgia, S.

Carolina." I have seen specimens from localities in Florida,

Oklahoma, Missouri, New Hampshire, Connecticut, and

Wisconsin. If communis is a synonym, then this species also

ranges into the Neotropics; however, neither Halstead

(1973:83) nor I have seen any specimens from south of the

United States.

Specimens examined. More than 180, of which 60 were

from Florida representing 13 collection records (for

complete data see Appendix).

Cathartosilvanus opaculus (LeConte), new combination

Silvanus opaculus LeConte 1856:78

Silvanus trivialis Grouvelle 1878:75, new synonym

Silvanus (Cathartosilvanus) trivialis, Grouvelle 1912a:340

Cathartosilvanus trivialis, Halstead 1973:84

Diagnosis. Length 1.7mm 2.1mm. Adults of this

species can be distinguished from those of imbellis by their

smaller size and strongly transverse pronotum.

67

Taxonomic notes. Halstead (in litt. and on

determination labels) examined the relevant types and

suggested this synonymy.

Biology. Halstead (1973:84) reported the following

associations: egg plants, pineapple, bananas, cedar and

mahogany logs, Primavera log, dried fruit of Musa textilis,

withered foliage of sugarcane and banana, in nest of weaver

bird, and at light. It occasionally is transported on

produce, but there are no records (to my knowledge) of it

being of economic importance.

Distribution. The type locality of opaculus is

"Colorado River, California" (LeConte 1856:78). Halstead

(1973:84) reported it from Central and South America south

to Brazil and Bolivia, as well as from Jamaica, Grenada,

Guadeloupe, and St. Vincent in the Caribbean, and from

California and Arizona. Thomas (1979) reported it from

southern Florida. I have seen specimens from additional

localities in Trinidad, U.S. Virgin Islands (St. Croix), and

the Dominican Republic. In Florida it has been collected

only in Dade County.

Specimens examined. More than 25, of which three were

from Florida, representing two collection records (for

complete data see Appendix).

68

Genus Silvanoprus Reitter

Silvanoprus Reitter 1911:45

Type species. Silvanus fagi Guerin-Meneville [by

monotypy].

Diagnosis. The following combination of character

states will distinguish adults of this genus: tarsomere III

lobed; femoral lines closed (fig. 61); anterior angles of

pronotum acute (fig. 8).

Taxonomic notes. Reitter (1911:45) described this genus

for Silvanus fagi, which does not have produced temples, to

distinguish it from other European Silvanus which have

temples. Grouvelle (1912a:341) pointed out that the

development of the temples is variable in this genus and was

not a good character. He wrote that the most consistent

character is a lobed tarsomere III. Grouvelle (1912a:341-

342) transferred 11 additional species to Silvanoprus and

added a new one. He also divided the genus into two groups,

a fagi group of seven species in which the notosternal

suture of the prothorax ends at the lateral edge of the

prothorax near the anterior angles, and a scuticollis group

of six species in which the suture attains the anterior

margin of the prothorax.

Biology. Horion (1960:162) recorded fagi from various

European hardwoods and conifers, and Vogt (1967:92) recorded

it from wreaths in cemeteries.

Fig. 8. Silvanoprus scuticollis (Walker), habitus. Length
2.2mm 2.5mm.

Distribution. Except for one tropicopolitan species

(scuticollis) and one species introduced into North America

(angusticollis), this genus is restricted to the Old World.

Silvanoprus scuticollis (Walker)

(fig. 8, 61)

Silvanus scuticollis Walker 1859:53

Silvanoprus scuticollis, Grouvelle 1912a:342

Silvanus triangulus Reitter 1876:60; syn. by Grouvelle

1912a:342

Diagnosis. Length 2.2mm 2.5mm. The combination of

lobed tarsomere III, closed femoral lines (fig. 61), and

acute anterior pronotal angles (fig. 8) characterize the

adults of this species in the Florida fauna.

Taxonomic notes. This is a distinctive, widespread

species with only the one recorded synonym.

Biology. Nothing has been recorded about the biology of

this widespread species. Most of the specimens I have seen

were collected at ultraviolet light traps. One specimen from

St. Croix was collected in an ethanol-baited vane trap. I

have never collected it under bark, even in areas which

produced specimens from ultraviolet light traps.

Distribution. Hetschko (1930:64) recorded it from East

Africa, France, Guyana, Madagascar, East Indies, Ceylon,

Malay Archipelago, Japan, Sumatra, Antilles, Grenada,

Guadeloupe, and St. Vincent. Thomas (1979:358) recorded it

from Florida, Georgia, and North Carolina. I can add the

following U.S. localities: Alabama, Louisiana, and Oklahoma.

I have also seen specimens from U.S. Virgin Islands (St.

Croix), Costa Rica, Taiwan, Dominican Republic, Panama,

Brazil, and Jamaica.

Specimens examined. More than 85, of which 43 were from

Florida, representing 22 collection records (for complete

data see Appendix).

Genus Monanus Sharp

Monanus Sharp 1879:85

Emporius Ganglbauer 1899:578; syn. by Sharp and Scott

1908:429

Type species. Of Monanus, M. crenatus Sharp [by

monotypy]; of Emporius, Monotoma concinnula Walker [by

synonymy of Silvanus signatus von Frauenfeld, type by

monotypy].

Diagnosis. The combination of lobed tarsomere III,

closed femoral lines, denticulate, setose pronotal margins,

and elytra with transverse dark fascia characterizes this

genus in the Florida fauna.

Taxonomic notes. Sharp and Scott (1908:429) synonymized

Emporius Ganglbauer with Monanus Sharp. Grouvelle

(1912a:344) divided Monanus into two subgenera, Monanus

(sens. str.) and M. (Monanops), separating them on the basis

of body form, relative length of elytra, and length of

metasternum. He listed four species in the subgenus Monanops

and 17 in Monanus (sens. str.), of which three are now

considered synonyms of concinnulus. Recently, Pal (1981)

revised the Indian members of the genus, adding a new

species of M. (Monanops), and describing the larva of

concinnulus. He compared members of this genus with those of

Airaphilus Redtenbacher, which also is restricted to the Old

World.

Biology. Pal (1981:241) reviewed the biology of the

species of Monanus, citing earlier records of M. concinnulus

in the roots of Lonchocarpus in Peru and in stored grain

products in Nigeria, as well as recording Indian species

from "...haystack, leaf garbage and under...leaf sheath..."

Distribution. According to Pal (1981:241-242), the

genus Monanus is "...restricted to warm climatic zones, a

few species extend to warm temperate zones but none occur in

the cool temperate climate. It is unrepresented in Europe."

Only the tropicopolitan M. concinnulus occurs in the New

world.

73

Monanus concinnulua (Walker)

(fig. 9)

Monotoma concinnula Walker 1858:207

Monanus concinnulus, Grouvelle 1912a:371

Silvanus signatus von Frauenfeld 1867:438; syn. by Grouvelle

1908b:489

Cryptamorpha fasciatus Wollaston 1873:169; syn. by

Waterhouse 1876:122

Cathartus fascipennis Reitter 1878:129; syn. by Ganglbauer

1899:586

Diagnosis. Length 1.9-2.2mm. According to Grouvelle

(1912a:370-371) individuals of concinnulus can be

distinguished from those of other species of Monanus by

their poorly developed temples, elytra about two times

longer than wide, antennomeres IV-VIII quadrate, elytra with

transverse fascia.

Taxonomic notes. The wide distribution of this species

has resulted in it being described under several names. The

synonymy listed above was based on that by Pal (1981:247).

Biology. Little has been published on the biology of

this widespread species. I collected specimens on an old,

moldy palm frond in Port-au-Prince, Haiti; the only Florida

specimens I have seen were collected on Bambusa vulgaris.

Wolcott (1950:298) reported it from bananas, grapefruit, and

pomarrosa (Eugenia ambos).

Fig. 9. Monanus concinnulus (Walker), habitus. Length 1.9mm
2.2mm.

75

Distribution. Found throughout the warm areas of the

world. In the New World it has been recorded (Blackwelder

1945:421) from Mexico, Nicaragua, Puerto Rico, Guadeloupe,

St. Vincent, Mustique, and Grenada. I have seen specimens

from Taiwan, India, Jamaica, Belize, Panama, U.S. Virgin

Islands (St. Croix and St. John), Haiti, and Florida. It is

not known whether it is established in Florida.

Specimens examined. More than 55, of which two were

from Florida, with the following data: "Coral Gables

Fla."/"C.F. Dowling, Jr. coll. 6-III-59"/"at Bambusa

vulgaris".

Genus Ahasverus Gozis

Ahasverus Gozis 1881:cxxvii

Type species. Cryptophagus advena Waltl [by monotypy].

Diagnosis. Members of this genus can be distinguished

by the following combination of character states: tarsomere

III lobed, lateral margins of pronotum simple, femoral lines

open, body ovate, intercoxal process of sternum III rounded

anteriorly.

Taxonomic notes. Halstead (in litt.) concluded that

this is a polyphyletic genus, with rectus and related

species forming a group deserving of generic status. He is

preparing a revision of this group that will clarify the

supraspecific relationships.

Biology. Based on literature records and my collecting

experience, these are rarely, if ever, found under bark.

76

Only the biology of A. advena has been studied to any

extent, and it seems to be primarily fungivorous, even in

stored products.

Distribution. Except for the cosmopolitan stored

products pest A. advena (and occasionally excisus), members

of this genus are restricted to the Nearctic and Neotropical

regions. There are three described species in the United

States; all occur in Florida.

Key to the adults of Florida species of Ahasverus

1. Antennomere XI acuminate apically (fig. 63);

anterior pronotal angles strongly lobed; body

ovate, strongly convex dorsally.....advena (Waltl)

1'. Antennomere XI not acuminate apically (fig. 64);

anterior pronotal angles not strongly lobed; body

more parallel-sided; not strongly convex.......... 2

2(1'). Pronotum transverse, at apex clearly wider than at

base (fig. 10a) ...................rectus (LeConte)

2'. Pronotum quadrate, at base slightly broader than

at apex (fig. 10c).............longulus (Blatchley)

77

Ahasverus advena (Waltl)

(fig. 10b, 63)

Cryptophagus advena Waltl 1834:169

Ahasverus advena, Gozis 1881:cxxvii

Latridius musaeorum Ziegler 1844:270; syn. by Fauvel 1889:132

Silvanus guerini Allibert 1847:12; syn. by Ganglbauer

1899:589

Silvanus angustatus Lucas 1847:221; syn. by Ganglbauer

1899:132

Cryptophagus striatus Rouget 1876:ccvii; syn. by Reitter

1880:86

Diagnosis. Length 1.8-2.2mm. The ovate, convex body

shape, produced anterior pronotal angles (fig. 10b), and

acuminate antennomere XI (fig. 63) distinguish this species.

Taxonomic notes. Despite the distinctiveness of this

species, its wide distribution has led to numerous synonyms.

It is most similar to excisus, a Neotropical species

sometimes also found in stored products and which can be

distinguished from advena by the very deep emarginations

behind the anterior pronotal angles.

Biology. Corbett et al. (1937) studied the biology of

this species on copra in Malaya and found that it preferred

copra infested with the surface molds Penicillium glaucum

and Aspergillus sp., and they rarely completed development

on good copra. "It is practically entirely mycetophagous

and, after having consumed the surface moulds, has been

78

observed on several occasions leaving good quality copra"

(Corbett et al. 1937:85). Wolcott (1950:298) reported that

it had been intercepted in Puerto Rico in lima beans and

pigeon peas, as well as being collected in dry pods of Inga

laurina and under bark of a dead tree. Smith (1910:262)

recorded it in New Jersey as ". rare under bark; more

common in stored grain, fruit, nuts, etc., particularly such

that are spoiled." Brimley (1938:179) reported it in North

Carolina from ears of corn. Other records include: Leonard

(1928:381) in New York in dried pears from California;

Loding (1945:77) in Alabama in cereals; Blatchley (1910:563)

in Indiana in ". various articles of commerce,

especially damp flour, meal, rice, figs, beans, apples,

etc."; Hatch (1961:205) "in copra, in coffee beans from

Colombia; in flour mills and grain elevators; in aerial

trap; in grass pile; in moldy peas and oats; and in sacked

grain"; Kirk (1969, 1970) in South Carolina in cured ham,

woods trash, corn bin, and stored oats. I have collected it

at light and in a pile of decaying soybeans in a mixed

hardwood-pine forest.

Distribution. Virtually worldwide in stored products

and established in many regions. In the United States it has

been recorded from New Jersey (Smith 1910:262), North

Carolina (Brimley 1938:179), New York (Leonard 1928:381),

Alabama (1945:77), Indiana (Blatchley 1910:563), Washington,

Idaho, and Oregon (Hatch 1961:205), South Carolina (Kirk

Fig. 10. Ahasverus spp. 10a) A. rectus (LeConte), habitus.
Length 1.5mm 1.9mm; 10b) A. advena (Waltl),
prothorax; 10c) A. longulus (Blatchley), prothorax.

1969, 1970), Florida, southern California, and Arizona (Leng

1920:198). I have seen specimens from Florida, Kentucky,

Indiana, Iowa, Arizona, Texas, New York, New Hampshire,

Vermont, Ontario, Washington, Massachusetts, Ohio, Oklahoma,

Kansas, Maryland, Wisconsin, Arkansas, California, Georgia,

and Missouri, as well as Cuba and Thailand.

Specimens examined. More than 50, of which which 16

were from Florida, representing 10 collection records (for

complete data see Appendix).

Ahasverus longulus (Blatchley)

(fig. 10c)

Cathartus longulus Blatchley 1910:564

Ahasverus longulus, Hetschko 1930:67

Silvanus parviceps Casey 1916:136; syn. by Halstead 1973:40

Diagnosis. Length 2.3-2.5mm. The non-acuminate

antennomere XI, anterior pronotal angles not strongly

produced, and quadrate pronotum which is slightly broader

basally than apically (fig. lOc) distinguish adults of this

species.

Taxonomic notes. Except for the original descriptions

and the synonymy proposed by Halstead (1973:40) nothing has

been published on the taxonomy of this species.

Biology. Blatchley (1910:564) reported that the type(s)

had been sifted from the borders of a sphagnum marsh.

Nothing else has been published on its biology, but (based

on specimens I have examined) it appears to be associated

with moist habitats. The single Florida specimen was

collected in an ultraviolet light trap.

Distribution. The type locality of longulus is Starke

Co., Indiana; that of parviceps is New York. Thomas

(1979:358) recorded it from Florida. I have seen specimens

from Florida, Missouri, Ohio, and Ontario.

Specimens examined. Nine, of which one was from Florida

with the following data: "FLA., Alachua County Gainesville

4-VII-1978 Coll. M.C. Thomas"/"blacklight trap" (MCTC).

Ahasverus rectus (LeConte)

(fig. 10a, 64)

Silvanus rectus LeConte 1856:78

Ahasverus rectus, Grouvelle 1912a:382

Diagnosis. Length 1.5-1.9mm. Adults of this species

are similar in general habitus to those of longulus but can

be distinguished by their strongly transverse prothorax,

which is wider anteriorly than basally.

Taxonomic notes. There are no recorded synonyms for

this widespread, common species. The record from Puerto Rico

(Wolcott 1950:298) probably refers to Ahasverus plagiatus

Grouvelle, which is very similar in external structure with

rectus.

Biology. Little has been recorded on the biology of

this species. Loding (1945:77) reported it as "Bred from

corn." Kirk (1969, 1970) recorded it from woods trash. It is

commonly collected in ultraviolet light traps. I have

82

observed it in Florida around grass tufts on sandy soil, in

grass clippings, and under old squares of sod. I have seen

numerous specimens, both adults and larvae, from pitfall

traps in an orange grove.

Distribution. It ranges throughout the southern United

States, west to Arizona, and has been recorded from North

Carolina (Brimley 1938:179), Alabama (Loding 1945:77),

"Pennsylvania, Illinois and southward." (Blatchley

1910:564), Illinois, District of Columbia, Arizona, and

Florida (Leng 1920:199). I have seen specimens from Mexico

and the following U.S. states: Georgia, Alabama, Missouri,

Arizona, Texas, North Carolina, and Florida.

Specimens examined. More than 250, of which 228 were

from Florida representing 83 collection records (for

complete data see Appendix).

Genus Cathartus Reiche

Cathartus Reiche 1854:77

Type species. Silvanus quadricollis GuBrin-Mgneville

[by synonymy of Cathartus cassiae Reiche, type by monotypy].

Diagnosis. Members of this genus can be

distinguished by the following combination of character

states: tarsomere III lobed, pronotal margins simple,

femoral lines open, antennomere XI at its broadest narrower

than X, intercoxal process of sternum III pointed

anteriorly.

Taxonomic notes. Many species, not strictly congeneric

with quadricollis, have been assigned at one time or another

to this genus. Hetschko (1930:70-71) listed four species,

but since the group has never been completely revised it is

uncertain if the other species, one each from Australia, New

Guinea, and Central America, are properly assigned.

Cathartus quadricollis, at least, seems to be most closely

related to those of Ahasverus.

Biology. Only the biology of Cathartus quadricollis, a

cosmopolitan stored products pest, has been reported. It

seems to be especially associated with corn, both stored and

in the field, and in the wild, pods of Tamarindus indica,

in which I collected numerous specimens in southern Dade

County, Florida. Wolcott (1950:298) reported it "in

abundance" in tamarind pods in Puerto Rico. On several

occasions I have also collected it from under the bark of

Ouercus spp. It is often taken in ultraviolet light traps.

Ganglbauer (1899:588) reported specimens in Germany from

Havana cigars and from Cassia fistula. Kirk (1969, 1970)

reported it from corn in the field, in corn bin, corn mill

trash, pea-vine hay, and on plum foliage. Loding (1945:77)

reported it from corn meal. Aitken (1975:127) reported it

from maize, cocoa beans, and copra residue, and as a field

pest of corn in Nigeria and Jamaica.

Distribution. Cathartus quadricollis is almost

cosmopolitan in distribution. The other three species

84

currently assigned to Cathartus were described from New

Guinea, Australia, and Honduras.

Cathartus quadricollis Gugrin-Mgneville

(fig. 11, 62)

Silvanus quadricollis Guerin-Meneville 1829-44:198

Cathartus quadricollis, Ganglbauer 1899:587-588

Cathartus cassiae Reiche 1854:78; syn. by Ganglbauer

1899:587

Silvanus gemellatus Jacquelin du Val 1857:104; syn. by

Ganglbauer 1899:588

Diagnosis. Length 2.1-3.2mm. The characters given in

this section under the genus will distinguish individuals of

this species in Florida.

Taxonomic notes. The identity of this species was

disputed for many years, primarily because of the great

sexual dimorphism it exhibits and the fact that GuBrin's

type of quadricollis has been lost. Ganglbauer (1899:587-

588) proposed the above synonymy, which was adopted by

Hetschko (1930) for his world catalogue and by most modern

authorities. Grouvelle (1912a:373) accepted the synonymy of

cassiae with gemellatus but argued that Gu6rin's description

of quadricollis actually applied to LeConte's Silvanus

(=Cathartosilvanus) imbellis.

Biology. See this section under the genus.

Distribution. Hetschko (1930:70) recorded it merely as

cosmopolitan. Aitken (1975:127) recorded specimens

Fig. 11. Cathartus quadricollis Gu6rin-Maneville,
habitus. Length 2.1mm 3.2mm

originating in Mexico, Ghana, Ivory Coast, Nigeria, and

Borneo, as well as Jamaica and Alabama. He suggested that it

is native to the southeastern United States. Sharp

(1899:559) recorded it from Mexico, Guatemala, Nicaragua,

Panama, and Cuba. Wolcott (1950:298) reported it from Puerto

Rico. Casey (1884:73) recorded it from Florida, Georgia, and

New York. Loding (1945:77) from Alabama; Blatchley

(1910:563) from Pennsylvania and Georgia; Hatch (1961:205)

from Washington and Oregon; Leonard (1928:381) from New

York; Kirk (1969, 1970) from South Carolina. I have seen

specimens from Argentina, Bahamas, Bolivia, Brazil, British

Honduras, Dominican Republic, Ghana, Jamaica, Mexico,

Panama, Trinidad, and, from the United States: North

Carolina, Florida, Texas, California, and District of

Columbia. In Florida it ranges throughout the state.

Specimens examined. More than 170, of which 103 were

from Florida, representing 31 collection records (for

complete data see Appendix).

PASSANDRIDAE

Diagnosis. Members of this family can be distinguished

by the following combination of character states: genae

produced anteriorly as rounded plates which conceal

maxillae; gular sutures confluent; aedeagus inverted, with

articulated parameres; tarsal formula 5-5-5 in both sexes;

dorsal surface of head and prothorax with system of lines

and grooves as in Laemophloeidae; elytra with system of

elytra cells or remnants thereof; hind wing venation well-

developed.

Larval diagnosis. Body hypergastric, swollen; head and

abdominal segment IX small; mouthparts reduced,

articulations not evident on maxilla or labium; antennae

short, inconspicuous, not articulated; legs short, stout;

spiracles annular; urogomphi short, hook-like.

Taxonomic notes. This group is composed of genera long

recognized as distinct, although it usually has been treated

as a subfamily of Cucujidae. However, Crowson (1955:102)

recognized its distinctiveness and raised it to family rank.

Thomas (1984a:78) considered that Passandridae is more

closely related to Laemophloeidae than to Cucujidae and

included it in a monophyletic lineage including also

Propalticidae and Phalacridae. Hetschko (1930) listed eight

genera (in three subfamilies of the Cucujidae) and 114

	Title Page
	Acknowledgement
	Table of Contents
	List of Figures
	Abstract
	Introduction
	Materials and methods
	Key to the genera of cucujidae,...
	Silvanidae
	Passandridae
	Laemophloeidae
	Summary
	Appendix
	References
	Biographical sketch
	Back Matter

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