Coleoptera: Cleridae) of Florida

Home , Bark beetle, Chariessa, Chramesus, Ciidae, Cleridae, Clerinae

THE CHECKERED BEETLES (COLEOPTERA: CLERIDAE) OF FLORIDA

JOHN MOELLER LEAVENGOOD, JR.

A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2008

In loving memory of Kevin Thomsen, a dear friend

ACKNOWLEDGMENTS

For their shared interest in taxonomy and their inspiration I would like to thank James C.

Dunford, Marc A. Branham, James C. Wiley, John L. Foltz, and Paul M. Choate. I would like to thank my supervisory committee (Michael C. Thomas, Paul E. Skelley, and Amanda C. Hodges)

for their support and friendly criticism.

I would also like to thank my father, John M. Leavengood. He has been a financier of this project and he has served me enormously as a confidant and a friend. A lifetime seems far too short to truly repay him for all he has done for me—but I will try anyway.

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF FIGURES ...... 8

LIST OF ABBREVIATIONS ...... 12

ABSTRACT ...... 13

CHAPTER

1 INTRODUCTION ...... 15

Literature Review ...... 18 Materials and Methods ...... 20 Key to the Subfamilies of Cleridae ...... 22

2 CLERINAE ...... 25

Key to the Florida Genera of Clerinae ...... 25 Genus Enoclerus Gahan, 1910 ...... 25 Key to the Florida Species of Enoclerus ...... 26 Enoclerus angustus (LeConte), 1849 (Figs. 2-1, 2-2) ...... 27 Enoclerus ichneumoneus (Fabricius), 1777 (Figs. 2-3, 2-4) ...... 29 Enoclerus lunatus (Klug), 1842 (Figs. 2-5, 2-6) ...... 35 Enoclerus nigrifrons (Say), 1823 (Figs. 2-7, 2-8) ...... 39 Enoclerus nigripes (Say), 1823 (Figs. 2-9, 2-10) ...... 40 Enoclerus rosmarus (Say), 1823 (Figs. 2-11, 2-12) ...... 42 Genus Placopterus Wolcott, 1910 ...... 44 Placopterus subcostatus (Schaeffer), 1917 ...... 44 Placopterus thoracicus (Olivier), 1795 (Figs. 2-13, 2-14) ...... 45 Genus Priocera Kirby, 1818 ...... 47 Priocera castanea (Newman), 1838 (Figs. 2-15, 2-16) ...... 48 Genus Thanasimus Latrielle, 1806 ...... 50 Thanasimus dubius (Fabricius), 1777 (Figs. 2-17, 2-18) ...... 51 Genus Trichodes Herbst, 1792 ...... 54 Trichodes apivorus Germar, 1824 (Figs. 2-19, 2-20) ...... 55

3 ENOPLIINAE ...... 70

Key to the Florida Genera of Enopliinae ...... 70 Genus Chariessa Perty, 1830 ...... 70 Key to the Florida Species of Chariessa ...... 71 Chariessa floridana Schaeffer, 1917 (Figs. 3-1, 3-2) ...... 71 Chariessa pilosa Forster, 1771 (Figs. 3-3, 3-4) ...... 71 Genus Cregya LeConte, 1861 ...... 76 Key to the Florida Species of Cregya ...... 76 Cregya oculata (Say), 1835 (Figs. 3-5, 3-6) ...... 76 Cregya quadrinotata Spinola, 1844 (Figs. 3-7, 3-8) ...... 79 Genus Neorthopleura Barr, 1976 ...... 80 Neorthopleura thoracica (Say), 1823 (Figs. 3-9, 3-10) ...... 80 Genus Pelonium Spinola, 1844 ...... 87 Pelonium leucophaeum (Klug), 1842 (Figs. 3-11, 3-12) ...... 87

4 EPIPHLOEINAE ...... 98

Key to the Florida Genera and Species of Epiphloeinae ...... 98 Genus Madoniella Pic, 1935 ...... 98 Madoniella dislocatus (Say), 1825 (Figs. 4-1, 4-2) ...... 98 Genus Pyticeroides Kuwert, 1894 ...... 100 Pyticeroides laticornis (Say), 1835 (Figs. 4-3, 4-4) ...... 101

5 HYDNOCERINAE ...... 106

Key to the Florida Genera of Hydnocerinae ...... 106 Genus Isohydnocera Chapin, 1917 ...... 106 Isohydnocera curtipennis (Newman), 1840 (Figs. 5-1, 5-2) ...... 106 Genus Phyllobaenus Dejean, 1833 ...... 108 Key to the Florida Species of Phyllobaenus ...... 110 Phyllobaenus humeralis (Say), 1823 (Figs. 5-3, 5-4) ...... 111 Phyllobaenus pallipennis (Say), 1825 (Figs. 5-5, 5-6) ...... 112 Phyllobaenus pubescens (LeConte), 1849 (Figs. 5-7, 5-8) ...... 114 Phyllobaenus rufipes (Newman), 1840 (Figs. 5-9, 5-10) ...... 116 Phyllobaenus suturalis (Klug), 1842 (Figs. 5-11, 5-12) ...... 117 Phyllobaenus unifasciatus (Say), 1825 (Figs. 5-13, 5-14) ...... 118 Phyllobaenus verticalis Say, 1835 (Figs. 5-15, 5-16) ...... 120

6 KORYNETINAE ...... 132

Genus Necrobia Olivier, 1795 ...... 132 Key to the Florida Species of Necrobia ...... 132 Necrobia ruficollis Fabricius, 1775 (Figs. 6-1, 6-2) ...... 133 Necrobia rufipes Degeer, 1775 (Figs. 6-3, 6-4) ...... 133 Necrobia violacea (Linnaeus), 1758 (Figs. 6-5, 6-6) ...... 137

7 TARSOSTENINAE ...... 142

Genus Tarsostenus Spinola, 1844 ...... 142 Tarsostenus univittatus (Rossi), 1792 (Figs. 7-1, 7-2) ...... 142

8 THANEROCLERINAE ...... 146

Key to the Florida Genera and Species of Thaneroclerinae ...... 146 Genus Ababa Casey, 1897 ...... 146 Ababa tantilla (LeConte), 1865 (Figs. 8-1, 8-2) ...... 147 Genus Thaneroclerus Lefebvre, 1838 ...... 148 Thaneroclerus buquet (Lefebvre), 1835 (Figs. 8-3, 8-4) ...... 149

9 TILLINAE ...... 153

Key to the Florida Genera of Tillinae ...... 153 Genus Callotillus Wolcott, 1911 ...... 153 Callotillus eburneocinctus Wolcott, 1911 (Figs. 9-1, 9-2) ...... 153 Genus Cymatodera Gray, 1832 ...... 155 Key to the Florida Species of Cymatodera ...... 156 Cymatodera bicolor (Say), 1825 (Figs. 9-3, 9-4) ...... 157 Cymatodera floridana Barr, 1972 (Figs. 9-5, 9-6) ...... 160 Cymatodera inornata (Say), 1835 (Figs. 9-7, 9-8) ...... 161 Cymatodera undulata (Say), 1825 (Figs. 9-9, 9-10) ...... 163 Cymatodera wolcotti Barr, 1950 (Figs. 9-11, 9-12) ...... 165 Genus Cymatoderella Barr, 1962 ...... 168 Cymatoderella collaris (Spinola), 1844 (Figs. 9-13, 9-14) ...... 169 Genus Lecontella Wolcott and Chapin, 1918 ...... 170 Lecontella brunnea (Spinola), 1844 (Figs. 9-15, 9-16) ...... 171 Genus Monophylla Spinola, 1841 ...... 173 Monophylla terminata (Say) 1835 (Figs. 9-17, 9-18, 9-19, 9-20) ...... 174

LIST OF REFERENCES ...... 187

BIOGRAPHICAL SKETCH ...... 206

LIST OF FIGURES

Figure page

2-1 Enoclerus angustus ...... 60

2-2 Distribution map of Enoclerus angustus...... 60

2-3 Enoclerus ichneumoneus ...... 61

2-4 Distribution map of Enoclerus ichneumoneus ...... 61

2-5 Enoclerus lunatus ...... 62

2-6 Distribution map of Enoclerus lunatus ...... 62

2-7 Enoclerus nigrifrons ...... 63

2-8 Distribution map of Enoclerus nigrifrons ...... 63

2-9 Enoclerus nigripes ...... 64

2-10 Distribution map of Enoclerus nigripes ...... 64

2-11 Enoclerus rosmarus ...... 65

2-12 Distribution map of Enoclerus rosmarus ...... 65

2-13 Placopterus thoracicus ...... 66

2-14 Distribution map of Placopterus thoracicus ...... 66

2-15 Priocera castanea ...... 67

2-16 Distribution map of Priocera castanea ...... 67

2-17 Trichodes apivorus...... 69

2-18 Distribution map of Trichodes apivorus ...... 69

3-1 Chariessa floridana ...... 92

3-2 Distribution map of Chariessa floridana ...... 92

3-3 Chariessa pilosa...... 93

3-4 Distribution map of Chariessa pilosa ...... 93

3-5 Cregya oculata ...... 94

3-6 Distribution map of Cregya oculata ...... 94

3-7 Cregya quadrinotata ...... 95

3-8 Distribution map of Cregya quadrinotata ...... 95

3-9 Neorthopleura thoracica ...... 96

3-10 Distribution map of Neorthopleura thoracica ...... 96

3-11 Pelonium leucophaeum ...... 97

3-12 Distribution map of Pelonium leucophaeum ...... 97

4-1 Madoniella dislocatus ...... 104

4-2 Distribution map from Madoniella dislocatus ...... 104

4-3 Pyticeroides laticornis ...... 105

4-4 Distribution map of Pyticeroides laticornis ...... 105

5-1 Isohydnocera curtipennis ...... 124

5-2 Distribution map of Isohydnocera curtipennis ...... 124

5-3 Phyllobaenus humeralis ...... 125

5-4 Distribution map of Phyllobaenus humeralis ...... 125

5-5 Phyllobaenus pallipennis ...... 126

5-6 Distribution map of Phyllobaenus pallipennis ...... 126

5-7 Phyllobaenus pubescens ...... 127

5-8 Distribution map of Phyllobaenus pubescens ...... 127

5-9 Phyllobaenus rufipes ...... 128

5-10 Distribution map of Phyllobaenus rufipes ...... 128

5-11 Phyllobaenus suturalis ...... 129

5-12 Distribution map of Phyllobaenus suturalis ...... 129

5-13 Phyllobaenus unifasciatus ...... 130

5-14 Distribution map of Phyllobaenus unifasciatus ...... 130

5-15 Phyllobaenus verticalis ...... 131

5-16 Distribution map of Phyllobaenus verticalis ...... 131

6-1 Necrobia ruficollis ...... 139

6-2 Distribution map of Necrobia ruficollis ...... 139

6-3 Necrobia rufipes...... 140

6-4 Distribution map of Necrobia rufipes ...... 140

6-5 Necrobia violacea ...... 141

6-6 Distribution map of Necrobia violacea...... 141

7-1 Tarsostenus univittatus ...... 145

7-2 Distribution map of Tarsostenus univittatus ...... 145

8-1 Ababa tantilla...... 151

8-2 Distribution map of Ababa tantilla ...... 151

8-3 Thaneroclerus buquet ...... 152

8-4 Distribution map of Thaneroclerus buquet ...... 152

9-1 Callotillus eburneocinctus ...... 177

9-2 Distribution map of Callotillus eburneocinctus ...... 177

9-3 Cymatodera bicolor ...... 178

9-4 Distribution map of Cymatodera bicolor ...... 178

9-5 Cymatodera floridana ...... 179

9-6 Distribution map of Cymatodera floridana ...... 179

9-7 Cymatodera inornata ...... 180

9-8 Distribution map of Cymatodera inornata ...... 180

9-9 Cymatodera undulata...... 181

9-10 Distribution map of Cymatodera undulata ...... 181

9-11 Cymatodera wolcotti ...... 182

9-12 Distribution map of Cymatodera wolcotti ...... 182

9-13 Cymatoderella collaris...... 183

9-14 Distribution map of Cymatoderella collaris ...... 183

9-15 Lecontella brunnea ...... 184

9-16 Distribution map of Lecontella brunnea ...... 184

9-17 Monophylla terminata, female ...... 185

9-18 Monophylla terminata, male ...... 185

9-19 Monophylla terminata, male (left) and female (right) ...... 186

9-20 Distribution map of Monophylla terminata ...... 186

LIST OF ABBREVIATIONS

CAPS Cooperative Agricultural Pest Survey

FDACS-DPI Florida Department of Agriculture and Consumer Services, Division of Plant Industry

FSCA Florida State Collection of Arthropods, Gainesville, Florida

JMLC John M. Leavengood, Jr., private collection

RHTC Robert H. Turnbow, private collection

UCFC University of Central Florida collection

UF-ENY University of Florida, Department of Entomology and Nematology

Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science

THE CHECKERED BEETLES (COLEOPTERA: CLERIDAE) OF FLORIDA

John M. Leavengood, Jr.

December 2008

Chair: Michael C. Thomas Major: Entomology and Nematology The Florida components of the beetle family Cleridae, with 42 species in 22 genera were reviewed: Ababa tantilla (LeConte), Callotillus eburneocinctus Wolcott, Chariessa floridana

Schaeffer, C. pilosa Forster, Cregya oculata (Say), C. quadrinotata (Chevrolat), Cymatodera bicolor (Say), C. floridana Barr, C. inornata (Say), C. undulata (Say), C. wolcotti Barr,

Cymatoderella collaris (Spinola), Enoclerus angustus (LeConte), E. ichneumoneus (Fabricius),

E. lunatus (Klug), E. nigrifrons (Say), E. nigripes (Say), E. rosmarus (Say), Isohydnocera curtipennis (Newman), Lecontella brunnea (Spinola), Madoniella dislocatus (Say), Monophylla terminata (Say), Necrobia ruficollis (Fabricius), N. rufipes Degeer, N. violacea (Linnaeus),

Neorthopleura thoracica (Say), Pelonium leucophaeum Klug, Phyllobaenus humeralis (Say), P. pallipennis (Say), P. pubescens (LeConte), P. rufipes (Newman), P. suturalis (Klug), P. unifasciatus (Say), P. verticalis Say, Placopterus subcostatus (Schaeffer), P. thoracicus

(Olivier), Priocera castanea (Newman), Pyticeroides laticornis (Say), Tarsostenus univittatus

(Rossi), Thanasimus dubius (Fabricius), Thaneroclerus buquet (Lefebvre), and Trichodes apivorus (Germar). Distinguishing characteristics of each genus were discussed. Each species treatment includes diagnostic discussion, biology, and distribution sections followed by label

data and selected references. A habitus photograph and a distribution map are presented for nearly all species.

CHAPTER 1 INTRODUCTION

The group of coleopterous insects treated here as a full family, the Cleridae, historically

has been recognized as multiple taxa. This is a product of the significant morphological

variation within the family. Most species are medium-sized and brightly colored (often with an

aposematic pattern) red, black and orange. The few small, dull species (Thaneroclerinae) are

often associated with stored product pests and are widely distributed. The subfamilies are rather varied in appearance compared to other families of Coleoptera, but are characterized by wide heads, usually wider than the pronotum, the pronotum often rather cylindrical and usually narrower than the elytra, a 5-5-5 tarsal formula with some members apparently with only four tarsal segments, and the entire body covered with bristle-like hairs. The elytral punctures of most taxa become obsolete in the apical fourth and members of several subfamilies may have a faint, pale midelytral fascia.

The elaborate color patterns exhibited by the family earned them the common name “the checkered beetles”. However, not all species have patterns that are “checkered” nor do all species have color patterns at all, although rarely so. Members of all subfamilies for which there are natural history accounts are known to be entomophagous as immatures and as adults, at least in part. Exceptions are adults which may use pollen as an alternative source of protein. Most entomophagous taxa are predacious on bark beetles and other wood-boring beetles. The larvae of many species have been recorded as predators of Hymenoptera. The Korynetinae and

Tarsosteninae prey upon stored product pests and larval Hydnocerinae have been observed feeding on chalcidoid larvae in galls as immatures while being commonly encountered on flowers as adults.

The Clerinae, Enopliinae, and Epiphloeinae are commonly encountered on trees infested

with Scolytidae, Buprestidae, Cerambycidae and Lyctinae (Bostrichidae). The larvae and, in

some cases, the adults can be found in the galleries opportunistically feeding on all life stages of

the prey species. When not found at the site of a host plant infestation some clerids will be found

on flowers, in high grassy habitats, or running up and down the stems and branches of trees and

shrubs. Adding intrigue to the Cleridae is that “unless one happens to be at the right place, at the

right time, when adult emergence and predation on emerging lignicolous insects are in temporal

synchrony…these [beetles] are likely to be gathered one or two specimens at a time”. These are

the words of Weston Opitz (2004: 6), who was referring to the subfamily Epiphloeinae, and it

rings true for all but the most common clerid taxa.

Ant mimicry is considered to be extremely common in the Cleridae and can be observed,

in part, by their movement patterns which are very similar to that of an ant scurrying rapidly up

and down plant substrates and shifting directions erratically (Mawdsley, 1994: 115, 2000: 166).

Several species are considered to be velvet ant mimics and others are presumed to be a part of a

large coleopterous mimicry ring including the Lampyridae and Lycidae as generalized models

(Mawdsley, 2002: 166).

Links to plant hosts are not strongly species-specific. Many species have been reared from

numerous plant hosts. The hydnocerines, if any, are the most specialized in that they can

reproduce in galls and domatia (Sabrosky, 1934: 65; Krombein, 1999: 1). Otherwise, they seem to have no narrower plant preferences than their prey species. Cymatodera, among a few other taxa, have been reared from seemingly undisturbed larval cells of Hymenoptera.

Members of the Necrobia are pests in some circumstances, but most often are only present in significant numbers when feeding on another stored product pest species. However, they will

feed on shipments of dried meats and sometimes on animal pelts and the like. Otherwise, clerids are not pests. Some taxa are known to control forest pest populations and are considered economically important. Thanasimus dubius and T. formicarius are best known for this. In fact,

T. formicarius was imported by Hopkins (1899) to biologically control forest pests of pine and spruce.

Most species demonstrate broad distribution and habitat range in the eastern United States.

In fact, most of the species which occur in Florida also occur in twelve to more than twenty other states, with many reaching Canada or Mexico. Among the Florida fauna, only three species are known to be precinctive (= endemic): Callotillus eburneocinctus, Cymatodera floridana, and

Chariessa floridana. One species of questionable status, Phyllobaenus suturalis, is known from

Florida and Mexico. The Korynetinae and Tarsosteninae often match the distribution of their stored product prey species, which are often cosmopolitan.

The Cleridae have received little attention in the form of revisionary work, especially at the family or tribe levels. As such, for the purpose of this thesis eight subfamilies are recognized:

Clerinae, Enopliinae, Epiphloeinae, Hydnocerinae, Korynetinae, Tarsosteninae, Thaneroclerinae and Tillinae, as per Opitz (2002). Opitz (2007: 79) suggested that it takes years to accumulate enough material for a “reasonable understanding of intraspecific variation”. The few and far between specimens collected in lesser known regions further supports the need for patience in the advancement of cleridology. Barr (quoted by Opitz) explained “time and time again I get a shipment and almost always am surprised to find something totally unexpected in the form of one specimen, invariably never to see another one of the same kind”. As a result, some species that have stood described for over twenty years on the basis of a single tropical specimen have never been collected since (as per observations of museum material over time).

Literature Review

Early publications on Florida Cleridae were limited largely to species and genus descriptions and the casual mention of observations on behavior, prey species or host plants.

The first significant works on Cleridae by Spinola (1841, 1844) and Klug (1842), which included species accounts numbering in the hundreds. Later, as with most beetle taxa, much descriptive work by LeConte, Gorham, Horn, Melsheimer, and Chevrolat contributed to the known species through the end of the 1800’s.

Bøving and Champlain (1920) addressed the larvae of North American Cleridae. While there was no revisionary work, some of their morphological discussions defended the family as a stable taxon. Five unnamed subfamilies were suggested, some of which match Opitz’s (2002) classification, others of which simply combine taxa of similar body form. Barr (1962: 121), in a non-revisionary work, suggested only two subfamilies of Cleridae, those being the Clerinae, represented by the tribes Tillini, Phyllobaenini, Thaneroclerini and Clerini, and the Korynetinae, represented by the tribes Epiphloeini, Enopliini and Korynetini. Opitz (2002: 268) commented that the current assignments of supraspecific groups in Cleridae were highly subjective, often lacking sufficient morphological, molecular and zoogeographical bases. Opitz considered a classification in which all of the tribes listed by Barr (1962) are recognized as subfamilies, plus

Tarsosteninae, pending further work.

Estimates by Papp (1960) differ little from Gerstmeier (1999) and Opitz (2002) as to the number of Cleridae worldwide (about 4000, 291 of which occur in the United States). Currently, some questions remain unanswered even for the North American fauna, where some intraspecific variations lend curiosity to the potential sinking or addition of species.

The morphology and behavior of these beetles shows potential for mimicry. This is especially evident in those species which resemble Lycidae and Lampyridae (Chariessa,

Perilypus and Pyticeroides among others) (Opitz, 1997: 51). Many species demonstrate variations which could lead to one species being mistaken for many. While insects of most taxa have undergone much synonymy and revision in the 250 years of modern classification, those with intraspecific color pattern variation create additional confusion. As a result, clerids have experienced a flurry of new, and subsequently, refuted species, some being recognized as several species for extensive periods of time. This creates a problem to those tracing the history of a species through dated literature whose references weave a labyrinthine mess of synonymy.

Such specific epithets as lampyriformis, formicarius and mutillaecolor suggest authors’ observations of similarities between the named clerids and other invertebrates. Mawdsley (1994:

115) explored potential relationships between the Cleridae and their possible models. The range of their mimicry spans many other families of Coleoptera, aculeate Hymenoptera, and even isolated cases mimicking Zygaenidae (Lepidoptera) and flower-visiting Tachinidae (Diptera).

While many visual mimics also show behavioral mimicry, only three species of the family have yet to demonstrate chemical defense. Should more species be found to possess chemical defenses, then there would also be Műllerian, rather than only Batesian, mimics within the family (Mawdsley, 2002: 166).

Many of the Clerinae, Enopliinae and Epiphloeinae are predacious as adults and immatures, making them important in identifying wood-borer infestations. Some clerids are the principle predators of wood and bark boring beetles and these predacious habits make them a taxon of economic importance (St. George, 1924: 49). As such, the family is generally associated with woody plants. Most of these clerids are encountered under bark, in the galleries of wood- and cone-boring insects, in galls or on dead twigs and branches. Many species of

Enoclerus and Thanasimus are considered to play an important role in the population control of

bark beetles in coniferous forests. Some of the Tillinae and Clerinae infiltrate nests and are

predators of bees and wasps while others are not predacious at all as adults, being found on

flowers and apparently feeding only on pollen. The immatures of Trichodes complete their

development on the egg pods of grasshoppers or within the cells of bees and aculeate wasps

(Opitz, 2002: 268, Knull, 1951: 271).

Others, especially hydnocerines, have been recorded to behave as endoparasites in cynipid

galls, the pupal cells of moths, and have even been encountered in egg nests of periodical cicadas

(Russel and Stoetzel, 1991: 385). Hydnocerines have been found in galls and in the pith-cavity of larval cells of Xylocopinae in domatia in Sri Lanka (Krombein, 1999: 28). Lecontella and

Cymatodera have been found in larval cells of Eumeninae (Vespidae), Megachilidae, and

Sphecidae.

While many clerids are predators of bark beetles of all life stages, and are considered forest

insects, Mawdsley (2002: 15) was the first to find direct associations between these beetles and

prairie flora. When found in grasslands clerids should not be expected to be found on their host

plants, but rather a broad range of plants as many adults feed as generalist predators on small

beetles, wasps, aphids, true bugs and flies. Because many clerids have densely pubescent bodies

and consume pollen as a secondary (or even primary) source of protein, they also function as

pollinators.

Materials and Methods

The specimens examined for this thesis were obtained from the holdings of the FSCA,

UCFC, JMLC, and RHTC.

Additional data was rendered from an effort by the Cooperative Agricultural Pest Survey

(CAPS). The project was the Solid Wood Packing Material survey for Sirex noctilio

(Hymenoptera: Siricidae). The author was the primary diagnostician for much of the statewide

material, which included checkered beetles. Insect specimens were identified and discarded.

Species were recorded into a database currently accessible to any member of CAPS via the I-

drive. Some checkered beetle specimens were preserved and curated at a later date. These

specimens were added to the JMLC and are not listed as CAPS material. The data resulting from

uncurated checkered beetles was entered into the CAPS database. In each species treatment

under the section “specimens examined” such data were included even though no voucher

specimens exist.

Checkered beetles were identified using various literature (primarily original species

descriptions), various microscopes, and using material from the JMLC and FSCA as reference

collections. All images of checkered beetles were photographed using the AutoMontage™ system at the FDACS-DPI and UF-ENY. Images were then edited using PhotoShop®.

Distribution maps were made using MapInfo Pro Viewer™ and cropped in Microsoft Office

Picture Manager®. Images and distribution maps were then converted from TIFF to JPEG files in Microsoft® Paint, and inserted into the thesis document as figures at the end of the appropriate

chapter.

There is an introduction chapter. Each following chapter represents one of the eight

subfamilies. Because the subfamilies have never been revised, they are addressed in this thesis

in alphabetical order. Similarly, the tribes and genera have received very little attention and no

revised genus with more than one Florida species is represented in this thesis. As such, the

species within each genus and the genera within each subfamily are also presented in

alphabetical order (and not to be mistaken for phylogenetic position).

Each species treatment discusses distinguishing characteristics and similar taxa, gives an

account of known biological behavior, and provides a full distribution followed by a county-

based distribution for Florida. In the “biology” section, the last paragraph often includes unreferenced information such as flight period or plant associations rendered from the label data of the “specimens examined”. The “specimens examined” section lists label data verbatim for

Florida specimens only, although other specimens were actually examined. Under each genus treatment the “selected references” section lists only literature which addresses all species of the genus. For species treatments, all known literature pertaining to the species in question is listed.

At the end of each chapter are habitus photographs and Florida distribution maps for nearly all species. The counties of the distribution maps are not labeled. See Figure 1-1 for county locations in Florida. In species distributions, Miami-Dade County is equivalent to Dade County.

Dichotomous keys to subfamilies, genera and species were made by combining characteristics from previous literature and adding characteristics limited to the Florida fauna.

The bibliography was formatted using the publishing guidelines of the journal Insecta

Mundi.

Key to the Subfamilies of Cleridae

1. Tarsomeres I-IV of front legs compact; tarsal pulvilli absent; entire body dull brown, patternless ...... Thaneroclerinae

1’. Tarsomeres I-IV of front legs not compact; tarsal pulvilli present; body variable ...... 2

2(1). Anterior coxal cavities closed behind ...... Tillinae

2’. Anterior coxal cavities open behind ...... 3

3(2). Eyes weakly emarginated; antennae short and stubby ...... Hydnocerinae

3’. Eyes deeply emarginated; antennae variable, but not short and stubby ...... 4

4(3). Tarsomeres III and IV subequal in length ...... Clerinae

4’. Tarsomere IV much smaller than I-III ...... 5

5(4). Antennal club as long or longer than remaining segments combined ...... 6

5’. Antennal club shorter than remaining segments combined ...... 7

6(5). Basal two antennal club segments somewhat apically forked, antler-like and elongate ...... Enopliinae

6’. Antennal club segments greatly dilated, quadrate to triangular in shape ...... Epiphloeinae

7(5). Body robust; elytra metallic blue or green, sometimes with orange humeri ...... Korynetinae

7’. Body elongate; elytra dark bluish-black with a pale midelytral fascia ...... Tarsosteninae

Figure 1-1. Map of Florida with counties labeled

CHAPTER 2 CLERINAE

Key to the Florida Genera of Clerinae

1. Pronotum bicolored in Florida species, orangish-yellow with a central black spot; elytra dark blue to bluish-black, but always unicolored in Florida species ...... Placopterus

1’. Pronotum unicolored in Florida species, reddish to black, sometimes with anterior region slightly paler than posterior region, but of the same general color; elytra variable, but never unicolored in Florida species ...... 2

2(1). Apical three antennomeres serrate, forming a club; elytra lacking a transverse white fascia or lunula at the middle or just posterior to the middle, instead with two pairs of yellowish midelytral maculae, two per elytron, the posterior pair being closer together than the more laterally placed anterior pair in Florida species ...... Priocera castanea (Fig. 2-15)

2’. Apical three antennomeres forming a club; elytra with or without a transverse white fascia or lunula at the middle or just posterior to the middle, never with two pairs of yellowish elytral maculae in Florida species ...... 3

3(2). Pronotum black, densely covered with ferrugineous hairs; elytra with alternating orangish- yellowish and blackish-purplish fasciae, lacking a transverse white fascia or lunula at the middle or just posterior to the middle ...... Trichodes apivorus (Fig. 2-19)

3’. Pronotum reddish, lacking dense ferrugineous hairs; elytra with or without a transverse white fascia or lunula at the middle or just posterior to the middle, never with the color pattern described above in Florida species ...... 4

4(3). Pronotum with deep, transverse subapical groove, joined by a median depressed line; third and fourth metatarsomeres not conspicuously dilated; elytra with humeri red, followed by alternating black and whitish fasciae ...... Thanasimus dubius (Fig. 2-17)

4’. Pronotum with at most a shallow, transverse, subapical groove; without a median depressed line; at least third and fourth metatarsomeres dilated; elytral with humeri red, following fasciae as above or not ...... Enoclerus

Genus Enoclerus Gahan, 1910

This is the largest clerid genus in the world, with about 200 species in the Western

Hemisphere (Mawdsley, 2001: 459). Six species occur in Florida.

The genus Enoclerus is medium-sized, elongate, and convex, and are distinguished by distinctly emarginate, finely granulate eyes, slender maxillary palpi, dilated labial palpi, a three- segmented antennal club which is rather abruptly enlarged, the pronotum with a shallow, transverse subapical groove lacking a median line joining it, tarsi that are apparently four- segmented, third and fourth segments of metatarsi dilated, and broadly dentate claws.

The species of Enoclerus can be diagnosed using coloration. Often, the differences in color are significant, such as having a black or pinkish-red abdomen or the presence or absence of a colored band in the anterior third of the elytra. In the presence of a reference collection, specimens of this genus probably need not be keyed out.

The adults and immatures of Enoclerus are predacious on bark beetles, weevils and other wood-borers. They have been associated with pine, spruce, elm, cedar, white oak, hickory, sweet gum, maple, peach and other hardwoods. These beetles may be observed moving along branches of vegetation with movement patterns similar to ants, particularly those of the genus

Camponotus (Hymenoptera: Formicidae). Rifkind (1997: 298) suggested that E. insidiosus

(Gorham) is a mimic of a particular sympatric ant, C. sericeiventris Guerin, in Costa Rica.

While E. insidiosus is not a North American species, it shares a general color pattern with many of its congeners. The midelytral pale fascia found in many species has darker coloration anterior and posterior to the midelytral region, creating a “wasp-waist” appearance.

Selected references: Fabricius, 1775: 157; Jacquelin du Val, 1860: 196; Gahan, 1910: 62; LeConte, 1849: 19; Horn, 1880: 150; LeConte and Horn, 1883: 218; Horn, 1885: 154; Wickham, 1895: 248; Wolcott, 1910b: 357, 1910c: 852, 1911: 118; Bøving and Champlain, 1920: 630; Bradley, 1930: 106; Bøving and Craighead, 1931a: 57; Wolcott, 1947: 78; Dillon and Dillon, 1972: 278; Barr, 1976a: 17; Mawdsley, 2001: 459; Opitz, 2002: 277.

Key to the Florida Species of Enoclerus

1. Each elytron with one crescent-shaped creamy white slightly elevated lunula located behind the middle, the punctation of the lunula being much less coarse and dense than the remaining

elytral regions, the pubescence on the lunula no more dense than the remaining elytral regions ...... Enoclerus lunatus (Fig. 2-5)

1’. Each elytron with a pale, unelevated fascia at the middle or behind the middle, with pubescence much more dense than on the basal third of the elytra ...... 2

2 (1). Elytra with a broad band of pinkish red across the middle third of the elytra, with an incomplete black fascia at its basal end and a complete black fascia at its apical end; integument of the basal fourth of the elytra distinctly darker than that of the band at the middle third ...... Enoclerus ichneumoneus (Fig. 2-3)

2’. Elytra lacking a black fascia in the basal fourth; integument in the basal third of the elytra with only one distinct reddish color ...... 3

3 (2). Abdomen red ...... 4

3’. Abdomen black ...... 5

4 (3). Elytra with a pale, often distinctly arched fascia at about the middle, followed by a much broader black fascia occupying one-fourth of the elytral length ...... Enoclerus nigripes (Fig. 2-9)

4’. Elytra with a pale fascia just behind the middle, the fascia broader marginally, narrowing towards the middle as the elytra meet, followed by a narrower black fascia ...... Enoclerus angustus (Fig. 2-1)

5 (3). Elytra with a midelytral pale fascia, anteriorly bordered by a black, somewhat incomplete fascia ...... Enoclerus rosmarus (Fig. 2-11)

5’. Elytra with a midelytral pale fascia, anteriorly bordered by no fascia ...... Enoclerus nigrifrons (Fig. 2-7)

Enoclerus angustus (LeConte), 1849 (Figs. 2-1, 2-2)

= Clerus rosmarus var. virginiensis Schaeffer, 1917

Barr (1976a: 20) clarified the status of this species from E. rosmarus var. virginiensis, which should not be considered a synonym of E. rosmarus.

Enoclerus angustus has blackish to reddish legs and is nearly entirely red to reddish-pink.

At the middle third of the elytra there are three adjacent transverse fasciae, two black fasciae with a white fascia between them. The white fascia is broadest marginally and narrows towards the middle as the elytra meet. The rest of the elytra is red to reddish-pink, occasionally with

slightly paler integument apically. Enoclerus angustus and E. nigripes are similar to E. rosmarus, which is distinguished by its black abdomen and a much broader black fascia posterior to the white fascia.

Biology: Adults have been collected in Florida from March to May. They have been collected in Malaise traps.

Distribution: Barr (1975: 10) recorded this species from Florida and Virginia. Papp

(1960: 84) stated that it occurs in the southeastern United States. Before 1950, any distributions

recorded for E. rosmarus may have included both E. rosmarus and E. angustus.

In Florida Enoclerus angustus has been recorded from Alachua, Duvall, Jefferson, Levy,

and Union Counties.

Specimens examined (16): FLORIDA: Alachua County: 2mi north of LaCrosse, J and E

Turf Farm, Santa Fe River flood plain, malaise trap, 10-V-1996, G.J. Steck and B.D. Sutton (1,

FSCA); 4mi north of LaCrosse: J and E Ranch, 5-V-1989, L.R. Davis, Jr. (1, FSCA); 9mi

northwest of Gainesville on SR 232, malaise trap, 5-V-1977, H.N. Greenbaum (1, FSCA);

Gainesville, 26-III-1948, E.D. McRae, 21-IV-1930, G.E. Merrill, 24-V-1964, R.E. White (3,

FSCA); SE Gainesville, Kincaid Rd., old field edge, xeric oak habitat, 6 meter Malaise trap, 1-8-

V-1999, B.D. Sutton (1, FSCA); 23-V-1969, M.L. May (1, FSCA); Duval County: Jacksonville,

8-V-1970, C.F. Zeiger (1, FSCA); Jefferson County: Monticello, 11-IV-1994 (1, FSCA); Levy

County: 24-III-1959, H.V. Weems, Jr. (4, FSCA); Union County: 1km south of Lake Butler on

Highway 121, 27-IV-1989, C.W. Mills, III (1, FSCA); Highway 241 at Santa Fe River, 16-IV-

1989, C.W. Mills, III (1, FSCA).

Selected references: LeConte, 1849: 21; Lohde, 1900: 46; Schaeffer, 1917: 131; Papp, 1960: 84; Barr, 1975: 10, 1976a: 20; Peck and Thomas, 1998: 85.

Enoclerus ichneumoneus (Fabricius), 1777 (Figs. 2-3, 2-4)

= Clerus rufus Olivier, 1791 = Enoclerus knabi Wolcott, 1910

Enoclerus ichneumoneus is a large, reddish-pink clerid with thoracic sternites ranging from

entirely red to entirely black, often with both colors present. The basal fourth of the elytra is

reddish, posteriorly margined by an incomplete transverse black fascia. The middle half is

reddish-pink, but of a paler shade than the basal fourth. The apical fourth follows with a black

fascia, then a white fascia, and a black elytral apex. The head and pronotum are densely, finely

punctate. The elytra are coarsely punctate basally, the punctures smaller apically.

Enoclerus lunatus is similar, but has an elevated, crescent-shaped midelytral fascia

margined with black. Another species, E. muttkowski, is nearly identical to E. ichneumoneus, but

it occurs in the northeastern states and does not pose a diagnostic problem in the southeast.

Bøving and Champlain (1920: 602) described the larvae and compared them to some related taxa.

Biology: According to Knull (1951: 298), adults of this species may be found on infested trees in the summer months and hibernating under bark in the winter months. They have been collected from Rhus spp. (Anacardiaceae), shagbark hickory, sweetgum, maple, white oak, Acer spp. (Sapindaceae), and red cedar infested with Pityophthorus sp., Scolytus quadrispinosus Say,

Phloeosinus sp. (Coleoptera: Curculionidae: Scolytinae), and Cerambycidae (Coleoptera)

(Bøving and Champlain, 1920: 633; Knull, 1951: 298). One overwintering event witnessed in

Pennsylvania found about twenty beetles packed together beneath thick bark (Bøving and

Champlain, 1920: 633).

Mawdsley (1999: 41) suggested that E. ichneumoneus and E. muttkowski (Wolcott), two

species from different lineages of Enoclerus, have converged to similar color patterns which may

mimic a mutillid wasp.

Specimens have been collected in Florida in all months except for January. Adults have

been collected at ultraviolet light, mercury vapor light, and black light, in Steiner Traps and in

Malaise traps. It has also been collected in both ethanol-baited and turpentine/frontalin-baited

Lindgren funnel traps. It has been collected on Citrus mitis Blanco (Rutaceae).

Distribution: Alabama, Arizona, Arkansas, California, Delaware, District of Columbia,

Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Maryland, Missouri, Mississippi, New Jersey,

New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas,

Utah, Virginia, West Virginia, Wisconsin and Ontario (Wolcott, 1909: 98; Wickham and

Wolcott, 1912: 57; Bøving and Champlain, 1920: 634; Wolcott, 1947: 79; Papp, 1960: 83; Barr,

1975: 7). Knull (1951: 298) recorded this species in Ohio from Cincinnati, which is on the

Kentucky border. Mawdsley (1999: 42) listed its distribution from Pennsylvania, south along the

Appalachians and foothills to Florida, and west to Wisconsin, Illinois and Kansas. This

distribution rules out the northeastern states included in the distribution of E. ichneumoneus

listed by many authors who often include ranges up to Canada and northern New York.

Mawdsley suggested that such northeastern are the result of confusing E. ichneumoneus with E.

muttkowski. Mawdsley included a table of characters to more easily separate the two species.

In Florida Enoclerus ichneumoneus has been recorded from Alachua, Brevard, Citrus,

Dixie, Duval, Flagler, Gadsden, Hardee, Hernando, Highlands, Hillsborough, Indian River,

Jackson, Lafayette, Lake, Leon, Liberty, Manatee, Martin, Miami-Dade, Marion, Nassau,

Orange, Pasco, Pinellas, Polk, Putnam, Saint Lucie, Seminole, Union, and Volusia Counties.

Specimens examined (206): FLORIDA: Alachua County: 29.36 N, 82.22 W, 28-VIII-

1992, J. Pickering (1, FSCA); 2mi northwest of Gainesville, 2-VII-1974, J.B. Heppner (1,

FSCA); Austin Carey Forest, flight trap, 21-IV-1976, G.B. Fairchild and Roberts (1, FSCA);

Gainesville, ethanol-baited Lindgren funnel trap, 14-IV-1989, T.H. Atkinson (6, FSCA);

Gainesville, Turpentine/frontalin-baited Lindgren funnel trap, X-2004, J.L. Foltz and J.M.

Leavengood, Jr. (1, JMLC); Gainesville, 19-VI-1968, L. O'Berry, 2-III-1972, G.B. Fairchild, 31-

V-1969, H.V. Weems, III, 4-VII-1960, H.V. Weems, Jr., 8-XII-1968, C.V. Weems, 1-VII-1966,

P.E. Frierson, 16-X-1967, C.R. Artaud, 7-IX-1986, P.E. Skelley, 21-X-1977, W.H. Whitcomb,

28-X-1978, 22-XI-1961, F.W. Mead, 8-VI-1982, L.A. Hetrick, 9-XI-1980, L. Darling, 25-X-

1961, [anon.], VI-2003, VII-2004, XII-2003, VII-2006, J.M. Leavengood, Jr., 5-IV-1973, H.

Greenbaum, 7-VI-1979, B. Gregory, Jr., 3-VI-1969, J.G. May, 6-IX-1990, M.C. Thomas, 14-

VIII-1979, H. Collins, 14-X-1980, R.E. Brown, 9-XI-1984, C.M. Stevens (20, FSCA; 5, JMLC);

Gainesville, Doyle Conner building, blacklight trap, 4-V-1986, 9-V-1976, 24-VI-1986, F.W.

Mead, malaise trap, 25-II-1982, [anon.], ultraviolet light trap, 9-VII-1990, F.W. Mead (6,

FSCA); Gainesville, Doyle Conner building, 25-III-1991, L.A. Stange, 13-VI-1991, F.W. Mead

(2, FSCA); Gainesville, NW 42nd Terr. and 34th Pl., ethanol-baited Lindgren funnel trap, X-

2005, XI-2005, J.L. Foltz and J.M. Leavengood, Jr. (2, JMLC); Gainesville, NW 42nd Terr. and

34th Pl., Turpentine/frontalin-baited Lindgren funnel trap, V-2005, VI-2005, 1-XII-2003, J.L.

Foltz and J.M. Leavengood, Jr., VI-2000, XI-2000, J.L. Foltz (5, JMLC); Gainesville, University of Florida Horticulture Experiment Station, 31-III-1976, G.B. Fairchild, flight trap, 20-III-1976,

G.B. Fairchild (2, FSCA); Gainesville, University of Florida, Surge/Natural Area Drive,

Turpentine/frontalin-baited Lindgren funnel trap, IV-1998, J.L. Foltz (2, JMLC); Gainesville,

610 NW 54th Terrace, 1-III-1995, L.A. Stange (1, FSCA); Gainesville: SW 20th Avenue, 12-III-

1991, C.W. Mills, III (1, FSCA); Highway 236, 2km west of Highway 241, 19-III-1999, 20-III-

1999, C.W. Mills, III (2, FSCA); Highway 241 and Highway 236, 26-III-1989, C.W. Mills, III

(1, FSCA); NW Gainesville, San Felasco State Park, 29-IV-1948, E.D. McRae, beating foliage,

2-VI-2001, E. Nearns (2, JMLC; 1, FSCA); Santa Fe River at Route 441, under bark, VII-1989,

P.E. Skelley (1, FSCA); SW Gainesville, I-75 and SW 20th, 11-II-1996, E. Nearns (1, JMLC);

SW Gainesville, near Tower Rd. and I-75, 3-IV-1995, E. Nearns (1, JMLC); 20-X-1946, (1,

FSCA); 23-IV-1970, M.L. May (1, FSCA); Brevard County: Cocoa, 17-III-1982, F.A. Smith (1,

FSCA); Merritt Island, ex. Quercus virginiana, 28-II-1985, A. Jones (1, FSCA); Titusville, SR

405, Enchanted Forest Sanctuary, White Trail, xeric oak hammock, Malaise Trap, 7-21-III-2001,

21-III-4-IV-2001, 4-18-IV-2001, 2-16-V-2001, 16-30-V-2001, 30-V-13-VI-2001, 27-VI-11-VII-

2001, 11-25-VII-2001, 1-15-VIII-2000, 31-VIII-14-IX-2000, 16-31-X-2000, 25-XI-14-XII-2000,

P. Russell, Z. Prusak and S.M. Fullerton (25, UCFC); Union Park, urban yard, 18-X-2000, A.

Hepburn (1, UCFC); University of Central Florida, MacKay Tract, Orlando, Sawgrass Marsh, red maple, Malaise Trap, 26-IV-1999, P. Russell and S.M. Fullerton (1, UCFC); University of

Central Florida, Orlando, LLP-Sand Pine, Turkey Oak, Malaise Trap, 2-V-1993, 17-VII-1997,

S.M. Fullerton (2, UCFC); University of Central Florida, Orlando, Longleaf Pine-Saw Palmetto,

Malaise Trap, 30-X-1997, S.M. Fullerton (1, UCFC); Citrus County: Homosassa, 15-IX-1975,

H.V. Weems, Jr. (1, FSCA); Dixie County: 4.2 miles north of Old Town, 15-V-1979, R.

Turnbow (1, RHTC); 4mi north of Old Town, 18-V-1979, R. Turnbow (1, FSCA); junction of highways 349 and 351, 30-V-1983, R. Turnbow (1, RHTC); Duval County: Jacksonville, ethanol-baited Lindgren funnel trap, 19-III-2007, 9-IV-2007, 30-IV-2007, 18-V-2007, 7-VI-

2007, 29-VI-2007, A. Johnson (10, CAPS); Mayport, 21-IX-1963, F.J. Santana, (1, FSCA); SE

Jacksonville, ethanol-baited Lindgren funnel trap, VI-2007, J.M. Leavengood, Jr. (1, JMLC);

Flagler County: Korona, 31-X-1977, J.N. Pott (1, FSCA); Gadsden County: Quincy, at blacklight, 6-IX-1971, (1, FSCA); Hardee County: Lemon Grove, Steiner tap, 3-VI-1964, R.H.

Rhodes (1, FSCA); Hernando County: Withlacoochee State Forest, Richloam Tract, Goat Road,

10-IV-1991, M.C. Thomas, 3-IV-1998, R. Morris (2, FSCA); Withlacoochee Tucker Hill, 25-V-

1994, J. Meeker (1, FSCA); Highlands County: Highlands Hammock State Park, 14-IV-1990, R.

Turnbow, 14-IV-1990, M.C. Thomas (1, RHTC; 1, FSCA); Hillsborough County: Lithia,

ethanol-baited Lindgren funnel trap, III-2007, J.M. Leavengood, Jr. (2, JMLC); Ruskin, ex.

Citrus sinensis, 21-X-1977, D.C. Chancey (1, FSCA); Tampa, ethanol-baited Lindgren funnel trap, III-2007, J.M. Leavengood, Jr. (1, JMLC); Indian River County: Vero Beach, ex. ripe

Morus sp. fruit, 23-III-1990, K. Hibbard (1, FSCA); Jackson County: Caverns State Park, 7-XII-

1957, H.V. Weems, Jr., 22-IV-1972, H. Greenbaum, 3-5-V-2008, M.C. and S. Thomas, 27-IV-

1986, R. Turnbow (6, FSCA; 4, RHTC); Lafayette County: 0.5 miles west of Branford, 29-V-

1988, R. Turnbow (1, RHTC); Lake County: Grand Island, ex. Citrus sp., 12-XI-1982, N.L.

Morrison (1, FSCA); Okahumpka, 23-V-1981, H.L. Bentley (1, FSCA); Leon County: Tall

Timbers Research Station, 6-IV-1985, R. Turnbow (1, RHTC); Liberty County: 14.8 miles north of Sumatra, 7-V-1988, R. Turnbow (1, RHTC); Apalachicola National Forest, ethanol-baited

Lindgren funnel trap, V-2007, J.M. Leavengood, Jr. (1, JMLC); Torreya State Park, beating, 22-

V-2004, E. Nearns, 3-V-2003, 6-V-1989, 7-V-1989, R. Turnbow, 16-V-1968, H.V. Weems, Jr.,

5-V-1989, M.C. Thomas (1, JMLC; 3, FSCA; 2, RHTC); Torreya State Park, malaise trap, 15-

VI-1974, H.V. Weems, Jr. and C.R. Artaud (1, FSCA); Manatee County: Palmetto, 5-X-1991,

L.D. Cunningham (1, FSCA); Rubonia, ex. Citrus mitis, 21-X-1977, D.C. Chancey (1, FSCA);

Terra Ceia, ex. guinea grass, 20-XI-1986, H. Gillis (1, FSCA); Steiner tap, 1-XI-1963, D.C.

Chancey (1, FSCA); Marion County: McIntosh, ethanol-baited Lindgren funnel trap, IV-2007,

J.M. Leavengood, Jr. (1, JMLC); Ocala, ethanol-baited Lindgren funnel trap, IV-2007, J.M.

Leavengood, Jr. (2, JMLC); Ocala National Forest, ethanol-baited Lindgren funnel trap, III-

2007, J.M. Leavengood, Jr. (1, JMLC); Miami-Dade County: Dade City, Steiner tap, 27-III-

1963, J.H. Grantham (1, FSCA); Nassau County: Fernandina Beach, Jackson trap, 29-III-1982, J.

Garbark (1, FSCA); Fort Clinch State Park, 5-V-1988, P.E. Skelley (1, FSCA); Orange County:

Apopka, alpha-beta pinene-baited Lindgren funnel trap, 28-X-1963, C.J. Musgrove (1, FSCA);

Orlando, alpha-beta pinene-baited Lindgren funnel trap, 29-III-2007, 27-IV-2007, 4-VI-2007,

29-XI-2006, B. Saunders (11, CAPS); Orlando, ethanol-baited Lindgren funnel trap, 29-III-2007,

28-VIII-2006, 30-XI-2006, 11-X-2006, B. Saunders (14, CAPS; 2, JMLC); Tangerine, 23-IX-

1982, D.C. Phelps (1, FSCA); Pasco County: New Port Richey, 12-VI-1980, L. Ippolite (1,

FSCA); Pinellas County: Saint Petersburg, ethanol-baited Lindgren funnel trap, VI-2007, J.M.

Leavengood, Jr. (1, JMLC); Putnam County: Crescent City, ethanol-baited Lindgren funnel trap,

III-2004, J.L. Foltz, J.M. Leavengood, Jr., J. White (1, JMLC); Saint Lucie County: Bluefield,

McPhail trap, 3-XI-1982, K. Hibbard (1, FSCA); Bluefield, 2-XII-1987, K. Campbell (1, FSCA);

Fort Pierce, 30-X-1985, K. Hibbard (1, FSCA); Seminole County: Lake Mary, 12-VI-1962, C.O.

Youtsey (1, FSCA); Sanford, Steiner trap, 17-V-1966, G.W. Desin (1, FSCA); 26-VII-1962,

G.W. Desin (1, FSCA); Union County: Highway 241 at Santa Fe River, 20-IV-1986, 10-XI-

1985, C.W. Mills, III (2, FSCA); Volusia County: Daytona Beach, VII-1960, W. Rosenberg (1,

FSCA); Holly Hill, 23-IX-1970, J.N. Pott (1, FSCA); Ormond Beach, ex. Citrus sp., 20-IV-1987,

J. Beckwith (1, FSCA); [county unknown]: New Zion, Steiner tap, 23-IX-1969, W.C. Rhodes (1,

FSCA).

Selected references: Fabricius, 1777: 230; Olivier, 1791: 13; LeConte, 1849: 20; Drury, 1879: 171; Schwarz, 1890: 165; Wickham, 1895: 250; Lohde, 1900: 43; Felt, 1906: 502; Wolcott, 1909: 98, 1910a: 321, 1910c: 853; Wickham and Wolcott, 1912: 57; Bøving and Champlain, 1920: 600, 602, 633; Wolcott, 1922: 73; Bøving and Craighead, 1931b: 276;

Beal and Massey, 1945: 69; Wolcott, 1947: 79; Barr, 1950: 62; Craighead, 1950b: 201; Knull, 1951: 297, pl. 8, fig. 29; Papp, 1960: 83; Barr, 1975: 7; Peck and Thomas, 1998: 85; Mawdsley, 1999: 41; Opitz, 2002: figs. 50, 54, 55, 56; Leavengood, 2008: (in press).

Enoclerus lunatus (Klug), 1842 (Figs. 2-5, 2-6)

= Clerus bicolor Melsheimer, 1846

Enoclerus lunatus is distinguished by the crescent-shaped, elevated midelytral lunula margined with black on each elytron just posterior of the middle. It is large and its integument is mostly red. The legs, mouthparts, antennae, thoracic sternites and abdomen are reddish to reddish-black. The head and pronotum are densely, finely punctate, and the elytra are coarsely punctate basally, the punctures smaller apically.

Enoclerus ichneumoneus is similar but has no elevated lunula. Possibly causing some confusion, E. nigripes has a pale arched fascia with black on either side just behind the middle of the elytra. However, the arched band in E. nigripes is densely pubescent and not elevated.

Biology: Enoclerus lunatus has been recorded on peach trees attacked by Scolytus rugulosus (Muller) (Coleoptera: Curculionidae: Scolytinae) in Georgia. It has also been observed on flowers (Bøving and Champlain, 1920: 634).

Adults have been collected in Florida during all months except for November. Specimens have been collected at lights, in McPhail traps and Malaise traps. Specimens have been collected on Cassia fasciculata (Michaux) Greene, Cassia bicapsularis Linnaeus blooms (Fabaceae), blooms of Castanea alnifolia (Linnaeus) Miller, Quercus virginiana Miller (Fagaceae), Citrus aurantium (= nobilis and paradisi) Linnaeus (Rutaceae), Ocimum micranthum Miller

(Lamiaceae), and Ceanothus microphyllus Michaux (Rhamnaceae). Adults have been reared from Metopium toxiferum (Linnaeus) Krug and Urban (Anacardiaceae) and Coccoloba diversifolia Jacquin (Polygonaceae). One specimen was collected in the nest of Solenopsis

geminata (Fabricius) (Hymenoptera: Formicidae). Label data did not suggest whether or not the

specimen of E. lunatus was collected alive.

Distribution: Alabama, Delaware, Florida, Georgia, Kansas, Maryland, Missouri, New

Jersey, New York, North Carolina, Pennsylvania, South Carolina, Texas and Virginia (Wolcott,

1909: 97, 1910b: 359, 1947: 80; Papp, 1960: 83; Barr, 1975: 7; Peck and Thomas, 1998: 85).

Wickham and Wolcott (1912: 58) recorded it from Riley County, Kansas, which is within 50

miles of Nebraska, and Orchard, Alabama, which is within 20 miles of Mississippi. Knull (1951:

299) suggested that it should occur in Ohio.

In Florida Enoclerus lunatus has been recorded from Alachua, Brevard, Clay, Collier,

Columbia, Dixie, Highlands, Indian River, Jefferson, Leon, Levy, Liberty, Marion, Miami-Dade,

Monroe, Orange, Palm Beach, Polk, Putnam, Saint Lucie, Seminole, Union, and Volusia

Counties. Wickham and Wolcott (1912: 58) recorded E. lunatus from Estero and Tampa, which are in Lee and Hillsborough Counties, respectively.

Specimens examined (189): FLORIDA: Alachua County: 1 mile south of Highway 26,

VI-1972, J. Macchi (1, FSCA); 2 miles north of Gainesville, ramp trap, 23-V-1974, H. Davis and

W. Jetter (1, FSCA); 5.5 miles west of Gainesville, Castlegate Mobile Home Park, L.R. Davis,

Jr., on Cassia fasciculata, L.R. Davis, Jr. (9, FSCA); Gainesville, on oak, 24-III-1953, B.K.

Dozier, 20-IV-1924, T.H. Hubbell, 7-V-1968, R.E. Woodruff, on blooms of Castanea alnifolia,

15-V-1968, 18-V-1968, R.E. Woodruff, VI-2003, J.M. Leavengood, Jr. (5, FSCA; 1, JMLC);

Brevard County: Malabar, Malabar Road, Malabar Scrub Sanctuary, fire unit 16, xeric oak scrub,

Malaise Trap, 22-VII-3-VIII-2001, P. Russell, Z. Prusak and S.M. Fullerton (1, UCFC);

Titusville, SR 405, Enchanted Forest Sanctuary, White Trail, xeric oak hammock, Malaise Trap,

21-III-4-IV-2001, 4-18-IV-2001, P. Russell, Z. Prusak and S.M. Fullerton (4, UCFC); Clay

County: Camp Crystal, 19-V-1962, H.V. Weems, Jr. (1, FSCA); Collier County: Naples, on

Citrus sinensis, 3-V-1995, M. Brodie (1, FSCA); Columbia County: 0.4 miles east of 41 on

Hammock Road, 30-IV-1978, R. Turnbow (1, RHTC); Dixie County: 3.5 miles north of Old

Town, on Route 349, 3-V-1979, M.C. Thomas (1, FSCA); 3.7 miles north of Old Town, 20-V-

1978, R. Turnbow (1, RHTC); 4 miles north of Old Town, 18-V-1979, R. Turnbow (1, RHTC);

Old Town, 20-V-1978, G.B. Edwards (1, FSCA); Highlands County: 2 miles south of Sebring,

15-IV-1990, M.C. Thomas (1, FSCA); Archbold Biological Station, insect flight trap, 23-V-

1978, 30-V-1978, H.V. Weems, Jr. and Klein, L.K., insect flight trap, 15-IV-1980, H.V. Weems,

Jr. and F.E. Lohrer, insect flight trap, 13-V-1979, 27-V-1979, H.V. Weems, Jr. and S. Halkin, insect flight trap, 15-VI-1979, H.V. Weems, Jr. and T. Weber (6, FSCA); Archbold Biological

Station, malaise trap, 14-IV-1980, 20-IV-1980, 22-IV-1980, 23-IV-1980, 26-IV-1980, 28-IV-

1980, 3-V-1980, 4-V-1980, L.L. Lampert, Jr., malaise trap, 11-IV-1991, 14-IV-1990, L.R.

Davis, Jr., malaise trap, 1-V-1978, H.V. Weems, Jr. (11, FSCA); Archbold Biological Station,

14-IV-1978, 21-IV-1979, 23-IV-1979, 1-V-1979, L.L. Lampert, Jr. (4, FSCA); Highlands

Hammock State Park, at light, 30-III-2002, M. Hrabovsky (2, FSCA); Lake Placid, on Citrus nobilis, 24-III-1950, Medsaer (1, FSCA); Sebring, fossil sand dunes, 20-III-1975, H.V. Weems,

Jr. (1, FSCA); Indian River County: 2.5 miles southeast of Wabasso, 16-V-1986, R. Turnbow (1,

RHTC); Vero Beach, 18-IV-1977, R. Carter (1, FSCA); Jefferson County: in nest of Solenopsis geminata, 3-VI-1958, C.L. Alligood (1, FSCA); Leon County: 4 miles west of Tallahassee, on

Highway 20, 30-IV-1985, C.W. O'Brien (1, FSCA); Levy County: 3.8 miles southwest of

Archer, malaise trap in rosemary and turkey oak sandhill, 15-V-1988, 21-V-1988, 19-VI-1988,

P.E. Skelley (5, FSCA); 5 miles north of Otter Creek, 13-V-1979, E. Giesbert (1, FSCA); Liberty

County: 4 miles north of Torreya State Park, 3-V-1980, M.C. Thomas (1, FSCA); Torreya State

Park, malaise trap, 17-V-1968, H.V. Weems, Jr. (1, FSCA); Torreya State Park, 3-V-2003, R.

Turnbow (1, RHTC); Marion County: north of Romeo, 18-IV-1982, M.C. Thomas (1, FSCA);

Ocala National Forest, Hopkins Prairie, insect flight trap, 11-V-1979, G.B. Fairchild (2, FSCA);

Oklawaha River, on Route 316, at light, 5-V-2002, M. Hrabovsky (2, FSCA); Miami-Dade

County: 2 miles southwest of Florida City, 29-V-1983, R. Turnbow (1, RHTC); 5 miles west of

Florida City, 21-V-1975, E. Giesbert (3, FSCA); Coral Gables, 13-VIII-1946, F. Hurst, on Citrus paradisi, 17-V-1932, B.F. Adams¸ X-1949, [anon.] (3, FSCA); Everglades National Park, 18-V-

1975, E. Giesbert (1, FSCA); Fuch's Hammock, near Homestead, insect flight trap, 24-V-1979,

T.S. Dickel and H.V. Weems, Jr. (1, FSCA); Homestead, 7-IV-1933, L.S. Light, Jr. (1, FSCA);

Kendall, 6-XII-1967, R.W. Swanson (1, FSCA); Miami, 11-VI-1965, B.K. Dozier (1, FSCA);

Richmond, 10-IV-1948 (3, FSCA); IV-1957, D. Thornton, IV-1948, 3-VI-1950, [anon.], V-1954,

W.T. Burch, 29-X-1955, D.R. Paulson (5, FSCA); Monroe County: Big Pine Key, 3-V-1958,

D.R. Paulson, on Ocimum micranthum, 10-IV-1959, H.V. Weems, Jr., 23-VI-1971, W.H. Pierce, emerged 2-VIII-1977, E. Giesbert (4, FSCA); Crawl Key, 19-VII-1974, B.K. Dozier (1, FSCA);

Grassy Key, emerged 14-VIII-1976, R. Turnbow (1, FSCA); Key Largo, 3-IV-1966, 2-V-1957,

H.V. Weems, Jr., 2-V-1958, [anon.], 14-V-1977, 15-V-1977, R. Turnbow, 16-V-1962, 10-VI-

1972, 29-VIII-1961, B.K. Dozier, 19-V-1975, E. Giesbert, 14-V-1977, 15-V-1977, 17-V-1979,

24-VI-1978, R. Turnbow, ex. Cocoloba diversifolia Jacq., emerged 4-X-1978, R. Turnbow (14,

FSCA; 10, RHTC); No Name Key, 14-I-1972, 17-II-1972, 13-IV-1971, W.H. Pierce, on Cassia

bicapsularis blooms, 24-II-1972, W.H. Pierce, 10-VI-1953, O.D. Link (11, FSCA); No Name

Key, ex. Metopium toxiferum L., emerged 19-VIII-1978, R. Turnbow (1, RHTC); north Key

Largo, 27-V-1965, R.M. Baranowski (2, FSCA); Stock Island, 5-X-1963, 5-IX-1961, H.V.

Weems, Jr. (2, FSCA); Tavernier, 20-X-1962, B.K. Dozier (1, FSCA); upper Key Largo, 15-V-

1969, 15-V-1979, E. Giesbert, 7-VI-1976, R. Turnbow (3, FSCA); 11-XII-1936, E.D. Barcus (1,

FSCA); Orange County: Orange County: FTU campus, Orlando, 16-IX-1977, S.M. Fullerton (1,

UCFC); Rk Spr Rn St Res, SPine/Oak Scrub, Malaise Trap, 24-V-1995, 27-IV-1995, 11-V-1995,

S.M. Fullerton (3, UCFC); Wekiwa Springs State Park, burn zone 43/31, S22 T20S R28E,

longleaf pine-turkey oak, Malaise Trap, 13-IV-2001, P. Russell and S.M. Fullerton (1, UCFC);

Winter Park, 29-IV-1936, 11-V-1931, 15-V-1931 (3, FSCA); on Citrus paradisi, 30-IV-1932,

H.K. Winter (1, FSCA); Palm Beach County: Boynton Beach, on Quercus virginiana, 10-XII-

1985, D. Leone (1, FSCA); McPhail trap, V-1936, O.D. Link (1, FSCA); Polk County: Lake

Marion Creek Estates, 4-IV-1998, 7-V-1998, R. Morris (4, FSCA); Putnam County: Grandin, 26-

VI-1992, L.R. Davis, Jr. (1, FSCA); 21-VI-1939, H. Hixson (1, FSCA); Saint Lucie County:

Indrio, 16-V-1986, R. Turnbow (1, RHTC); Seminole County: Econ. Wild. Area, scrub oak/saw palmetto burn site, Malaise Trap, 8-VII-2000, 10-VI-2000, 20-V-2000, 24-VI-2000, 26-VIII-

2000, T. Smith, P. Russell and S.M. Fullerton, 29-IV-2000, 6-V-2000, 27-V-2000, 17-VI-2000,

15-VII-2000, 30-VII-2000, T. Smith (9, UCFC; 15, FSCA); lower Wekiwa River State Preserve, burn zone, LW-5, S39 T19S R29E, longleaf pine-turkey oak, Malaise Trap, 2-IV-2001, P.

Russell and S.M. Fullerton (1, UCFC); Union County: on Highway 241, 1 kilometer north of the

Santa Fe River, 15-V-1987, C.W. Mills, III (1, FSCA); Volusia County: Cassadaga, on

Ceanothus microphyllus, 30-III-1962, H.V. Weems, Jr. (1, FSCA).

Selected references: Klug, 1842: 294; Spinola, 1844: 255, pl. 24, fig. 2; Melsheimer, 1846: 307; Hamilton, 1895: 335; Lohde, 1900: 43; Wolcott, 1909: 97, 1910b: 359; Wickham and Wolcott, 1912: 58; Bøving and Champlain, 1920: 634; Wolcott, 1947: 80; Knull, 1951: 299, pl. 9, fig. 34; Papp, 1960: 83; Ekis, 1975: 29; Barr, 1975: 7; Peck and Thomas, 1998: 85; Opitz, 2002: figs. 67, 70.

Enoclerus nigrifrons (Say), 1823 (Figs. 2-7, 2-8)

Enoclerus nigrifrons has a red head, pronotum, legs, antennae and mouthparts. The basal third of the elytra is red, followed by an arched, white midelytral fascia which is not anteriorly

margined with black. The remaining apical portion of the elytra is black except for an anteapical white spot, sometimes broadened to an incomplete fascia. Adults are 6 to 7 mm long.

Biology: Enoclerus nigrifrons has been recorded from southern cypress and pine infested with Scolytinae and other wood-borers (Coleoptera: Curculionidae) (Knull, 1951: 296). King and Fox (1970: 133) observed it attacking Ips sp. (Coleoptera: Curculionidae: Scolytinae) and trapped adults in “Ips-baited traps”.

In Florida an adult was collected in May. In Ohio adults have been collected from May to

July (Knull, 1951: 296).

Distribution: Alabama, District of Columbia, Florida, Indiana, Massachusetts, Nebraska,

Ohio, Pennsylvania, South Carolina, and Ontario (Wolcott, 1909: 97, 1947: 80; Knull, 1951:

296; Papp, 1960: 83; Barr, 1975: 9). Enoclerus nigrifrons is only known from Florida by a single specimen from Walton County.

Specimens examined (1): FLORIDA: Walton County: Highway 285 at Shoal River, 5-V-

1985, R. Turnbow (1, RHTC).

Selected references: Say, 1823: 190; LeConte, 1849: 23; Horn, 1885: 154; Wolcott, 1909: 97, 1910c: 853; Wickham and Wolcott, 1912: 55; Wolcott, 1922: 72; Beal and Massey, 1945: 87, 143; Wolcott, 1947: 80; Knull, 1951: 295; Papp, 1960: 83; King and Fox, 1970: 133; Barr, 1975: 9.

Enoclerus nigripes (Say), 1823 (Figs. 2-9, 2-10)

Enoclerus nigripes has a red abdomen, red thoracic sternites and black legs. The basal third of the elytra is red, followed by a black fascia, and then a pale, somewhat arched fascia.

The next fascia is black and very broad. The apex of the abdomen is pale. The head and pronotum are densely, finely punctate. The elytra are coarsely punctate basally, the punctures growing smaller moving apically. Adults are 5 to 7 mm long.

Enoclerus nigripes is most similar to E. rosmarus, which has a black abdomen and red legs. Enoclerus angustus and E. lunatus also share similarities with E. nigripes (see the discussions of E. angustus and E. lunatus for distinguishing characters).

King and Fox (1970: 133) observed specimens of E. nigripes (without subspecies or variation noted) which underwent a change in abdominal color from red to black, suggesting temporal dimorphism. Because one such beetle oviposited when it had both a red and black abdomen, the change was not attributed to sexual maturity.

Biology: Enoclerus nigripes has been recorded on spruce, American elm, white pine, and juniper infested with Hylurgopinus rufipes (Eichoff) (Coleoptera: Curculionidae: Scolytinae),

Pissodes strobi Peck (Coleoptera: Curculionidae), on which it feeds on adults, larvae and pupae

(Bøving and Champlain, 1920; 603; Pechuman, 1937: 8; Knull, 1951: 294). Other collection records include butternut, ash, mulberry and wild cherry (Chittenden, 1890: 154). It has been found in leaf litter in February in Ohio and is active from April to July (Knull, 1951: 294).

Mawdsley (1994: 119) considered it to be an ant mimic.

A single Florida specimen was collected from a log of Carpinus sp. (Betulaceae) in

February.

Distribution: Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Kansas,

Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, New Hampshire, New York,

North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Virginia, Wisconsin and Manitoba (Wolcott, 1909: 97, 1947: 80; Knull, 1951: 295; Papp, 1960: 83; King and Fox,

1970: 133; Barr, 1975: 9; Gosling, 1980: 69). Because many past authors offer the distribution of E. nigripes without differentiating the distribution of the various subspecies, many older reviews and checklists should be considered to address all forms of E. nigripes. All forms,

collectively, would expand the above listed distribution west to Arizona and California, and south to Mexico. Enoclerus nigripes has been recorded in Florida from Liberty County.

Specimens examined (1): FLORIDA: Liberty County: Torreya State Park, ex. dead

Carpinus log, 27-II-1998, P.E. Skelley (1, FSCA).

Selected references: Say, 1823: 191; Spinola, 1844: 264, pls. 23, 25, fig. 3, 4; LeConte, 1849: 24, 1859a: 122; Drury, 1879: 171; Horn, 1885: 154; Chittenden, 1890: 154; Lohde, 1900: 44; Felt, 1906: 666; Wolcott, 1909: 97, 1910c: 853; Wickham and Wolcott, 1912: 56; Bøving and Champlain, 1920: 603, 631; Wolcott, 1922: 71; Champlain and Knull, 1925: 113; Pechuman, 1937: 8; Hoffmann, 1940: 56; Beal and Massey, 1945: 78, 92; Wolcott, 1947: 80; Knull, 1951: 294, pl. 7, fig. 28; Papp, 1960: 83; King and Fox, 1970: 133; Dillon and Dillon, 1972: 278, pl. 28, fig. 11; Ekis, 1975: 30; Barr, 1975: 9; Gosling, 1980: 69; Mawdsley, 1994: 119; Peck and Thomas, 1998: 85; Mawdsley, 1999: 43.

Enoclerus rosmarus (Say), 1823 (Figs. 2-11, 2-12)

= Clerus oculatus Say, 1835

Enoclerus rosmarus has red legs and thoracic sternites and a black abdomen. The basal third of the elytra is reddish. At the middle third of the elytra there are three adjacent transverse fasciae, two black fasciae with a white fascia between them, and the apical third of the elytra is reddish. The posterior black fascia is much broader than the anterior one. The head is finely, sparsely punctate and the pronotum is finely, densely punctate. The elytra are subopaque and coarsely, densely punctate basally, irregularly arranged, the punctures growing smaller moving apically. Adults are 3.5 to 7.0 mm long.

This species resembles E. nigripes and E. angustus (see the discussions of E. nigripes and

E. angustus for distinguishing characters). Bøving and Champlain (1920: 603) described the larvae and compared them to some related taxa. However, it is uncertain as to whether they described the then undescribed E. angustus.

Biology: Knull (1951: 297) observed that E. rosmarus is common and abundant on flowers and weeds in May, June and July. Dillon and Dillon (1972: 279) frequently observed

adults on greater horseweed. Gosling (1980: 69) collected adults by beating dead sumac

branches. Mawdsley (2002b: 17) observed adults on flowers feeding on small braconids. In

captivity they readily consumed small flies and true bugs, and members of Curculionidae,

Mordellidae, Dermestidae and Aphidae. As such, Mawdsley considered it to be a generalist as an adult. It has been observed visiting Asclepias syriaca Linnaeus (Asclepiadaceae), Melilotus

officinalis (Linnaeus) Pallas (Fabaceae), Solidago sp., Conyza canadensis (Linnaeus) Cronquist

(Asteraceae), and Tradescantia virginiana Linnaeus (Commelinaceae).

Enoclerus rosmarus has been collected in Florida in April and May. Adults have been

reared from Solidago sp. (Asteraceae) and Rhus sp. (Anacardiaceae).

Distribution: Alabama, Arkansas, Delaware, District of Columbia, Florida, Georgia,

Illinois, Indiana, Iowa, Louisiana, Kansas, Kentucky, Maine, Maryland, Massachusetts,

Michigan, Mississippi, Missouri, Nebraska, New Jersey, New York, North Carolina, Ohio,

Oklahoma, Pennsylvania, Virginia, Wisconsin, and Ontario (Barr, 1975: 9; Gosling, 1980: 69;

Peck and Thomas, 1998: 85). It is possible that Barr (1975) based the distribution of previous

accounts published prior to 1950. Before 1950, any distributions recorded for E. rosmarus

should be considered to include both E. rosmarus and E. angustus. In Florida Enoclerus

rosmarus has been recorded from Jackson, Leon, and Liberty Counties.

Specimens examined (6): FLORIDA: Jackson County: Florida Caverns State Park, 6-V-

1968, G.H. Heinrich (1, FSCA); Leon County: Tall Timbers Research Station, 13-V-1968, G.H.

Heinrich (1, FSCA); Liberty County: Torreya State Park, 26-IV-1986, 5-V-1984, R. Turnbow (4,

RHTC).

Selected references: Say, 1823: 190, 1835: 163; Spinola, 1844: 267, pl. 26, fig. 1; LeConte, 1849: 21, 1859a: 121; Drury, 1879: 171; Lohde, 1900: 46; Wolcott, 1910c: 854; Schaeffer, 1917: 131; Bøving and Champlain, 1920: 600, 603, 631; Wickham and Wolcott, 1922: 56; Wolcott, 1927: 64, 1947: 81; Knull, 1951: 296, pl. 8, fig. 30; Papp, 1960: 84;

Dillon and Dillon, 1972: 278, pl. 29, fig. 9; Ekis, 1975: 32; Barr, 1975: 9; Gosling, 1980: 69; Peck and Thomas, 1998: 85; Mawdsley, 2002b: 17.

Genus Placopterus Wolcott, 1910

Placopterus is represented by four species throughout the eastern, central and southern

United States and Texas (Barr, 1962: 126).

Placopterus is recognized by emarginate and finely granulate eyes, emarginate labrum,

cylindrical and elongate apical maxillary palpi, dilated and securiform apical labial palpi, the

other segments elongate as well, and basally broadly dentate tarsal claws. They have oblong to

ovate bodies and submoniliform antennae, the last three segments forming a rather compact club,

the last segment of which being subtriangular and compressed apically. They have five

tarsomeres, apparently four.

Placopterus superficially resemble Enoclerus and are distinguished by the yellow

markings on the lateral pronotum, whereas all Florida Enoclerus have an entirely reddish

pronotum. Because specimens of only P. thoracicus were available for examination, no key is

presented for the genus.

Selected references: Chevrolat, 1876: 5; Wolcott, 1910b: 363; Leng, 1920: 149; Wolcott, 1947: 77; Barr, 1962: 126; Dillon and Dillon, 1972: 275; Opitz, 2002: 278.

Placopterus subcostatus (Schaeffer), 1917

Placopterus subcostatus is very similar to the more common P. thoracicus, but with dark blue elytra with somewhat irregularly elevated longitudinal lines. Adults are about 6 mm long.

Biology: Nothing is known of its biology.

Distribution: Wolcott (1947: 77) and Barr (1975: 6) recorded P. subcostatus from Texas and Florida. In Florida, P. subcostatus has been recorded from Alachua, Hernando, Highlands,

Levy, Liberty, Marion, Pasco, Putnam, Seminole, and Union Counties (Peck and Thomas, 1998:

85). Schaeffer (1917: 131) recorded it from Enterprise, Volusia County, Florida.

Specimens examined: No Florida specimens were available for examination. All FSCA specimens have been loaned out for a review of the genus.

Selected references: Schaeffer, 1917: 131; Wolcott, 1947: 77; Barr, 1975: 6; Peck and Thomas, 1998: 85.

Placopterus thoracicus (Olivier), 1795 (Figs. 2-13, 2-14)

= Thanasimus monilis Melsheimer, 1846 = Clerus ornaticollis LeConte, 1880

This species is distinguished by its dark blue to bluish-black somewhat shining elytra and yellow to orangish-red pronotum which has a central black spot. The head is finely, sparsely punctate. The shining, finely punctured pronotum has a transverse rounded depression at the apex and at the base. The elytra are densely, coarsely punctate. Adults are 5 to 7.5 mm long.

Biology: Placopterus thoracicus has been collected from butternut, willow, witch hazel, sumac, white oak, red maple, and hickory infested with Magdalis olyra (Herbst), Curculio sp.

(Coleoptera: Curculionidae), Lymantor decipiens LeConte, Scolytus rugulosus (Muller),

Micracis swainei Blackman, Chramesus icoriae LeConte (Coleoptera: Curculionidae:

Scolytinae), Melasis pectinicornis Melsheimer (Coleoptera: Eucnemidae), Anthaxia viridicornis

Say, Agrilus defectus LeConte (Coleoptera: Buprestidae), Pogonocherus parvulus LeConte,

Anelaphus parallelus (Newman) and Oncideres sp. (Coleoptera: Cerambycidae) (Knull, 1932:

42; 1934: 207, 1951: 291; Gosling, 1980: 67).

Placopterus thoracicus has been found overwintering in the pupal cells of Magdalis olyra

(Knull, 1951: 291). It is especially common on foliage in damp meadows (Dillon and Dillon,

1972: 275). Gosling (1980: 67) observed it at ultraviolet lights in hickory-oak habitats.

Foster and Barr (1972: 123) noted that it had been reared from Trypoxylon sp. and

Pemphredon lethifer Wagner (Hymenoptera: Sphecidae). Eliason and Potter (2000: 555) found larvae in dissected stem galls of Callirhytus cornigera (Osten Sacken) (Hymenoptera:

Cynipidae) on pin oak. It was unsure exactly what the larvae predated since many other insects occupied the galls.

Mawdsley (1994: 116) considered it to be a part of a tremendous coleopteran mimicry ring with Lampyridae, Cantheridae, Lycidae, Buprestidae, Meloidae, Ptilodactylidae, Elateridae, and the Cleridae Monophylla terminata, Pyticeroides laticornis and Chariessa pilosa.

Adults have been collected in Florida from February to May. In Ohio adults occur in May,

June and July (Knull, 1951: 291). Specimens have been collected at mercury vapor lights, ultraviolet lights and Malaise traps.

Distribution: Connecticut, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky,

Missouri, Nebraska, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee,

Texas, Virginia, Canada and Mexico (Bøving and Champlain, 1920: 630; Knull, 1951: 291;

Papp, 1960: 81; Foster and Barr, 1972: 123; Barr, 1975: 6; Mawdsley, 1999: 41; Eliason and

Potter, 2000: 555). Wolcott (1947: 77) recorded it from California. This record is most likely P.

thoracicus pallipes. In Florida P. thoracicus has been recorded from Alachua, Clay, Dixie,

Hernando, Highlands, Liberty, Marion, Okaloosa, Orange, Santa Rosa, Union, and Walton

Counties.

Specimens examined (28): FLORIDA: Alachua County: Archer, on Route 26, on oak,

11-IV-1992, F.W. Skillman, Jr. (1, FSCA); Gainesville, Malaise trap, 22-27-II-1975, H.N.

Greenbaum (1, FSCA); Gainesville, IV-2003, J.M. Leavengood, Jr. (1, JMLC); Clay County:

Goldhead Branch State Park, edge habitat sweeping, IV-2004, D. Serrano and J.M. Leavengood,

Jr. (1, JMLC); Dixie County: 3.4 miles north of Old Town, 11-IV-1980, R. Turnbow (2, RHTC);

Hernando County: Withlacoochee State Forest, 10-IV-1991, R. Turnbow (1, RHTC); Liberty

County: Apalachicola Bluffs and Ravines Preserve, 18-IV-1999, 3-V-2003 R. Turnbow (2,

RHTC); Torreya State Park, 2-IV-2005, J.M. Leavengood, Jr. and E. Nearns, 5-V-2007, J.M.

Leavengood, Jr. (2, JMLC); Marion County: Citra, NW 24th Avenue, at mercury vapor and ultraviolet light, 22-III-1993, 26-III-1993, 13-IV-1993, F.W. Skillman, Jr. (6, FSCA); Mosa

Bluff, 3 miles south of 314A, on oak, 7-IV-1992, F.W. Skillman, Jr. (3, FSCA); Okaloosa

County: Henderson Beach State Recreation area, 23-III-2003, R. Turnbow (1, RHTC); Orange

County: Walt Disney World, C-4 Stout Site, S 15, 16 T 24S R 27E, xeric oak/flatwoods, Malaise

Trap, 1-7-IV-1998, Z. Prusak and S.M. Fullerton (1, UCFC); Santa Rosa County: 1.7 miles north of Munson, 20-IV-1991, R. Turnbow (1, RHTC); 3.7 miles north of Munson, at mercury vapor and black light, 10-IV-1999, R. Turnbow (4, RHTC); Walton County: Defuniak Springs, 25-III-

1989, R. Turnbow (1, RHTC).

Selected references: Olivier, 1795: 18, pl. 2, fig. 22; Melsheimer, 1846: 307; LeConte, 1849: 23; Drury, 1879: 171; LeConte, 1880: 194; Chittenden, 1890: 154; Wolcott, 1910b: 362, 1910c: 854; Wickham and Wolcott, 1912: 55; Leng, 1920: 149; Bøving and Champlain, 1920: 630; Wolcott, 1922: 68; Blackman and Stage, 1924: 44; Knull, 1932: 42, 1934: 207; Blackwelder, 1939: 33; Wolcott, 1947: 77; Craighead, 1950: 203; Knull, 1951: 290, pl. 6, fig. 23; Papp, 1960: 81; Dillon and Dillon, 1972: 275, pl. 29, fig. 3; Foster and Barr, 1972: 123, 1975: 6; Gosling, 1980: 67; Mawdsley, 1994: 116; Peck and Thomas, 1998: 85; Mawdsley, 1999: 41; Eliason and Potter, 2000: 555; Opitz, 2002: figs. 59, 69, 120.

Genus Priocera Kirby, 1818

Priocera is represented by twelve species throughout North and Central America. Two additional species, P. pusilla Kirby and P. lecontei Wolcott, questionably recorded from

California and “North America”, respectively, may be synonyms (Barr, 1962: 125).

Priocera is distinguished by coarsely granulate eyes, thickened femora, serrate antennae which lack a club and elongate apical maxillary palpomeres.

Priocera resembles some species of Cymatodera and Enoclerus. Enoclerus can be separated by its three-segmented antennal club. The apical antennomere of Cymatodera is three to four times as long as it is wide (at middle) while that of Priocera is one and one-half to two

times the length and Priocera lacks any form of projection or tubercle on the sides of its

pronotum.

Selected references: Kirby, 1818: 389; LeConte, 1849: 17; Horn, 1880: 150; LeConte and Horn, 1883: 218; Wolcott, 1910c: 851; Bøving and Champlain, 1920: 619; Bradley, 1930: 106; Bøving and Craighead, 1931a: 56; Wolcott, 1947: 76; Barr, 1962: 125; Opitz, 2002: 277.

Priocera castanea (Newman), 1838 (Figs. 2-15, 2-16)

= Priocera rufescens Spinola, 1844 = Priocera maculata Ziegler, 1845 = Notoxus leprieurii Spinola, 1844

This clerine is mostly reddish-brown, including the basal third and apical fourth of the elytra. This middle portion of the elytra is black with two yellow maculae, one behind the middle and one in front of the middle positioned more laterally, on each elytron bordering the black and reddish-brown regions. The surface of the head is finely punctate. The pronotum is longer than wide with irregular elevated regions, basally constricted and with a transverse depression, and sparsely, irregularly punctate. The elytral surface has rows of coarse punctures which become obsolete apically. Adults are around 9 mm in length. Priocera castanea is similar in body form to members of Enoclerus and some Cymatodera (see the generic discussion of Priocera for distinguishing characters).

Biology: This nocturnal species has been collected on logs of white oak infested with larvae of Melittomma sp. (Coleoptera: Lymexylonidae), pine attacked by Dendroctonus valens

LeConte and pitch pine infested with Dendroctonus frontalis Zimmermann (Coleoptera:

Curculionidae: Scolytinae) (Bøving and Champlain, 1920: 628; Knull, 1951: 289). Gosling

(1980: 67) observed it at ultraviolet lights in oak-hickory habitats.

Adults have been collected in Florida from March through August and September.

Specimens have been collected at mercury vapor, ultraviolet and blacklights, under bark, in

Malaise traps and in ethanol-baited Lindgren funnel traps.

Distribution: Alabama, Connecticut, Florida, Georgia, Illinois, Indiana, Kansas, Maine,

Maryland, Michigan, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma,

Pennsylvania, South Carolina, Texas, Virginia and Ontario (Wickham and Wolcott, 1912: 54;

Wolcott, 1947: 76; Papp, 1960: 80; Barr, 1975: 5). Knull (1951: 289) recorded it in Ohio from

Scioto County, which is within 50 miles of West Virginia, and Cincinnati, which is on the

Kentucky border. Bøving and Champlain (1920: 628) recorded it from Arizona. Because no other species of Priocera were included in their work, it is suspected that this distributional note could be a product of misidentification since none of the cited checklists include such extreme western distribution. In Florida Priocera castanea has been recorded from Alachua, Clay,

Gadsden, Highlands, Liberty, Marion, Miami-Dade, Orange, Seminole, St. Lucie, and Volusia

Counties.

Specimens examined (48): FLORIDA: Alachua County: Gainesville, at light, 15-VII-

1995, M.C. Thomas, 8-IV-1980, 20-VIII-1978, M.C. Thomas (3, FSCA); Gainesville, Beville

Hts., blacklight trap, 19-III-1980, L.A. Stange (1, FSCA); Gainesville, Doyle Conner building, blacklight trap, 14-V-1985, F.W. Mead (1, FSCA); NW Gainesville, San Felasco State Park,

under bark, 15-IV-2002, E. Nearns and R. Morris (1, JMLC); Clay County: Goldhead Branch

State Park, 8-VI-1986, Cicero (5, FSCA); Gadsden County: Aspalaga Landing, at boat ramp, at

mercury vapor and black light, 5-V-2007, J.M. Leavengood, Jr. (1, JMLC); Highlands County:

Archbold Biological Research Station, 18-IV-1976, 5-V-1981, L.L. Lampert, Jr., 12-IV-1984,

N.M. Downie (5, FSCA); Archbold Biological Station, 5 miles south of Lake Placid and 1.8

miles south of SR 70/old SR 8, IV-2004, J.M. Leavengood, Jr., J.C. Dunford, K. Barbera (1,

JMLC); Liberty County: Torreya State Park, at blacklight, 21-23-V-1983, K.W. Vick, on a log,

15-VII-1987, P.E. Skelley, 7-V-1989, C.W. Mills, III, at mercury vapor and black light, 6-V-

1989, R. Turnbow, 4-6-V-2007, J.M. Leavengood, Jr., flood plain forest, at ultraviolet light, 24-

V-2003, E. Nearns and R. Morris (5, FSCA; 3, RHTC; 6, JMLC); Marion County: Ocala, ethanol-baited Lindgren funnel trap, IV-2007, J.M. Leavengood, Jr. (1, JMLC); Miami-Dade

County: Miami, ethanol-baited Lindgren funnel trap, VI-2007, J.M. Leavengood, Jr. (1, JMLC);

Orange County: University of Central Florida, Orlando, LLP-Sand Pine, Turkey Oak, Malaise

Trap, 30-V-1997, 8-V-1993, 17-V-1993, 21-VIII-1997, S.M. Fullerton (4, UCFC); University of

Central Florida, Orlando, LLP-Sand Pine, Turkey Oak, ultraviolet light trap, 27-III-1994, 10-V-

1994, 25-IV-1994, 7-IX-1994, 18-VII-1994, S.M. Fullerton (6, UCFC); Seminole County:

Oviedo, 1-V-1997, (1, UCFC); St. Lucie County: ethanol-baited Lindgren funnel trap, 30-IV-

2007, 25-V-2007 B. Saunders (2, CAPS); Volusia County: Daytona Beach, 1950, W. Rosenberg

(1, FSCA).

Selected references: Newman, 1838: 380; Spinola, 1844: 119, pl. 4, fig. 3; Ziegler, 1845: 268; LeConte, 1849: 17; Drury, 1879: 171; Hamilton, 1895: 335; Lohde, 1900: 24; Wolcott, 1910c: 851; Wickham and Wolcott, 1912: 54; Bøving and Champlain, 1920: 620, 628; Bøving and Craighead, 1931b: 276; Wolcott, 1947: 76; Craighead, 1950: 203; Knull, 1951: 289, pl. 5, fig. 19; Papp, 1960: 80; Ekis, 1975: 20; Barr, 1975: 5; Gosling, 1980: 67; Peck and Thomas, 1998: 86; Opitz, 2002: figs. 63, 73.

Genus Thanasimus Latrielle, 1806

Thanasimus is represented by five North American species which are widely distributed, generally favoring northern temperate climates (Opitz, 2002: 277).

Thanasimus is recognized by anteriorly emarginate, finely granulate eyes, loose, very gradually clavate three-segmented antennal club, the last segment of which is not apically truncate, filiform maxillary palpi with the apical segment slender and tapering apically, and the

labial palpi with segments slightly elongate, last segment compressed and elongate-triangular in shape. The tarsi are apparently four-segmented, slender, elongate, and the claws are broadly toothed. Thanasimus is distinguished from Enoclerus by a deep, transverse, subapical groove connected by a median depressed line on the pronotum.

Thanasimus formicarius was imported by Hopkins in the first attempt to biologically control a forest pest of spruce and pine (Hopkins, 1899: 1). Considerable research has been conducted on pheromonal interactions between Thanasimus, bark beetles and host tree chemical volatiles.

Mawdsley (1994: 119) considers many members of this genus to be ant mimics.

Selected references: Latrielle, 1806: 270; Schaeffer, 1777: pl. 137; Jacquelin, 1860: 196; LeConte, 1849: 19; Horn, 1880: 150; LeConte and Horn, 1883: 218; Wickham, 1895: 250; Wolcott, 1910c: 854; Bradley, 1930: 106; Bøving and Craighead, 1931a: 57; Wolcott, 1947: 77; Dillon and Dillon, 1972: 278; Opitz, 2002: 277.

Thanasimus dubius (Fabricius), 1777 (Figs. 2-17, 2-18)

= Thanasimus ruficeps Spinola, 1844

The head and pronotum are densely, finely punctate. The elytra are coarsely punctate basally, with the punctures becoming finer moving apically. The antennae and legs vary from red to black. The body is nearly entirely red, except for the elytra in which only the basal fifth is red. Immediately posterior to the basal region there is a black irregularly margined transverse fascia followed by additional alternating fascia of similar form of white, black and white. The apical portion of the elytra is black. Adults are 7 to 9 mm in length. Bøving and Champlain

(1920: 601) described the larvae and compared them to some related taxa.

Biology: Thanasimus dubius is an obvious predator of the southern pine beetle,

Dendroctonus frontalis Zimmerman (Coleoptera: Curculionidae: Scolytinae). Both adult and

larval stages of T. dubius are predacious on all life stages of bark beetles from egg to adult. The

clerid maintains an impressive level of “spatial and temporal coincidence with its prey, and

particularly so with the southern pine beetle populations” (Dixon and Payne, 1979: 178). It is

also a predator of many other bark beetles including Ips, Dendroctonus, Polygraphus

(Scolytinae), Pissodes (Coleoptera: Curculionidae), and other wood-borers in coniferous trees

(Bøving and Champlain, 1920: 629). Most such predation occurs in spruce, pine and elm trees

(Knull, 1951: 292). Mawdsley (1999: 40) noted that adult males and females, like those of many

other clerid species, frequently conceal themselves beneath bark or in bark crevices.

Adults have been collected in Florida in all months except for August and September.

Specimens have been collected in Malaise traps and Lindgren funnel traps baited with ethanol,

both turpentine and ethanol, and alpha-beta pinene. It has been collected on Pinus elliottii

Engelmann (Pinaceae).

Distribution: Alabama, Alaska, Colorado, Florida, Georgia, Indiana, Kansas, Louisiana,

Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Nebraska, New Jersey,

New Hampshire, New Mexico, New York, North Carolina, Pennsylvania, South Carolina,

Tennessee, Texas, Vermont, Virginia, Wisconsin, Ontario and Mexico (Wolcott, 1909: 96, 1947:

77; Papp, 1960: 82; Barr, 1975: 7). Wickham and Wolcott (1912: 54) recorded it from Chicopee,

Massachusetts, which is within 15 miles of Connecticut, and Mount Katahdin, Maine, which is

within 75 miles of Quebec and New Brunswick. Knull (1951: 292) suggested that it should

occur in Ohio. In Florida T. dubius has been recorded from Alachua, Baker, Bay, Duval,

Escambia, Leon, Liberty, Marion, Miami-Dade, Orange, and Santa Rosa Counties.

Specimens examined (153): FLORIDA: Alachua County: Gainesville, on Pinus, 19-X-

1917, E.W. Berger, 31-X-1932, [anon.], on Pinus elliottii, 2-I-1967, 8-X-1969, R.E. Woodruff,

IX-2003, XII-2003, J.M. Leavengood, Jr., 14-VII-2001, 16-VII-2001, T. Smith (6, FSCA; 5,

JMLC); Gainesville, frontalin and turpentine-baited Lindgren funnel trap, VI-1995, V-1995, J.L.

Foltz, 20-X-1994, E. Ash and D. Durgee (26, FSCA); Gainesville, Devil's Millhopper State Park,

Millhopper Drive, Turpentine/frontalin-baited Lindgren funnel trap, III-1999, XI-1999, XII-

1998, XII-1999, J.L. Foltz, II-2006, J.L. Foltz and J.M. Leavengood, Jr. (5, JMLC); Gainesville,

south side of Biven's Arm Lake, Lindgren funnel trap, 17-31-I-2003, P.E. Skelley (1, FSCA);

Gainesville, University of Florida, Surge/Natural Area Drive, Turpentine/frontalin-baited

Lindgren funnel trap, III-1999, IV-1998, J.L. Foltz, 17-XI-2003, 1-XII-2003, J.L. Foltz and J.M.

Leavengood, Jr. (4, JMLC); 15-XI-1941 (1, FSCA); Baker County: Glen Saint Mary, 10-III-

1958, E.W. Holder, Jr. (1, FSCA); Bay County: alpha-beta pinene-baited Lindgren funnel trap,

24-X-2006, 28-XI-2006, 19-XII-2006, C. Street (3, CAPS); ethanol-baited Lindgren funnel trap,

24-X-2006, C. Street (1, CAPS); Duval County: alpha-beta pinene-baited Lindgren funnel trap,

18-V-2007, A. Johnson (1, CAPS); Escambia County: alpha-beta pinene-baited Lindgren funnel trap, 5-I-2007, 24-I-2007, 13-III-2007, 11-X-2006, 19-XI-2006, 28-XI-2006, C. Street (41,

CAPS); Leon County: alpha-beta pinene-baited Lindgren funnel trap, 26-I-2007, 14-III-2007,

17-XI-2006, M. Bentley (18, CAPS); Liberty County: Apalachicola National Forest, ethanol- baited Lindgren funnel trap, V-2007, J.M. Leavengood, Jr. (1, JMLC); Marion County: Ocala

National Forest, ethanol-baited Lindgren funnel trap, III-2007, J.M. Leavengood, Jr. (3, JMLC);

Miami-Dade County: Miami, ethanol-baited Lindgren funnel trap, III-2007, V-2007, J.M.

Leavengood, Jr. (3, JMLC); Miami, alpha-beta pinene-baited Lindgren funnel trap, 16-III-2007,

R. Griffiths (1, CAPS); Orange County: RSRSR, T20S R29E S19, 17-VII-2001, Buffalo Tram.

Res. Staff (18, UCFC); Wekiwa Springs State Park, burn zone 43/31, S22 T20S R28E, longleaf

pine-turkey oak, Malaise Trap, 30-XII-2001, P. Russell and S.M. Fullerton (1, UCFC); Apopka,

on Mammillaria from Texas, 10-I-1985, C. Phelps (1, FSCA); Santa Rosa County: alpha-beta

pinene-baited Lindgren funnel trap, 5-I-2007, 26-X-2006, 9-XI-2006, 19-XI-2006, 15-XII-2006,

C. Street (12, CAPS).

Selected references: Fabricius, 1777: 229, 1801: 280; Spinola, 1844: 189, pl. 14, fig. 3; LeConte, 1849: 24; Hopkins, 1893a: 155, Hamilton, 1895: 335; Hopkins, 1899: 262; Lohde, 1900: 36; Wolcott, 1909: 96, 1910c: 854; Wickham and Wolcott, 1912: 54; Bøving and Champlain, 1920: 601, 628; Hoffmann, 1942: 7; Beal and Massey, 1945: 84, 86, 142, 143, 145; Wolcott, 1947: 77; Craighead, 1950: 201; Knull, 1951: 292, pl. 6, fig. 24; Papp, 1960: 82; Mignot and Anderson, 1969: 305; Dillon and Dillon, 1972: 278, pl. 28, fig. 13; Barr, 1975: 7; Dixon and Payne, 1979: 178; Gosling, 1980: 67; Peck and Thomas, 1998: 86; Mawdsley, 1999: 40; Opitz, 2002: figs. 62, 71.

Genus Trichodes Herbst, 1792

Trichodes is represented by eleven North American species of broad distribution (Barr,

1975: 11). The generic name means “shaggy”, most likely indicative of how the body is densely clothed with hairs (Dillon and Dillon, 1972: 279).

Most species of Trichodes exhibit similar behavior, being found primarily on flowers that are in bloom. Wickham and Wolcott (1912: 58) suggested Compositae and Umbelliferae as likely flower taxa on which one would find Trichodes. Their color patterns, behavior and biology make them likely mimics of wasps (Mawdsley, 1994: 120).

Trichodes is recognized by its deeply emarginate, finely granulate eyes, three-segmented triangular antennal club with the last segment apically truncate, subcylindrical maxillary palpi with the last segment longer and slightly wider those preceding, and strongly dilated labial palpi.

The body is very hairy and has a range of color patterns from yellow to metallic bluish-black.

The somewhat trigonal apical maxillary palpomere separates it from Enoclerus, which has a more cylindrical palpomere. Bøving and Champlain (1920: 611) described the larvae and compared them to some related taxa.

Selected references: Herbst, 1792: 154; Fabricius, 1801: 283; Latrielle, 1804: 150; Hope, 1840: 139; LeConte, 1849: 17; Lacordaire, 1857: 459; Crotch, 1870: 42; Horn, 1876b: 231, 1880: 150; LeConte and Horn, 1883: 218; Wolcott, 1910b: 367, 1910c: 851; Wickham and Wolcott, 1912: 58; Bøving and Champlain, 1920: 611, 635; Leng, 1920: 150; Bradley,

1930: 106; Balduf, 1935: 107; Wolcott, 1944b: 54; Bøving and Craighead, 1931a: 57; Wolcott, 1947: 81; Arnett, 1960: 599; Papp, 1960: 85; Barr, 1961: 109, 1962: 121; Ekis and Gupta, 1971: 60; Dillon and Dillon, 1972: 279; Barr, 1975: 11; Foster, 1976c: 1; Mawdsley, 1994: 120; Opitz, 2002: 277.

Trichodes apivorus Germar, 1824 (Figs. 2-19, 2-20)

= Trichodes trifasciatus Sturm, 1826 = Trichodes cribripennis Spinola, 1844 = Trichodes apivorus var. interruptus Wolcott, 1910 = Trichodes apivorus var. borealis Wolcott and Chapin, 1918

Trichodes apivorus is a large clerine with reddish to orangish-yellow occupying a median transverse band, an anteapical band, and the basal fifth of the elytra. The rest of the body and elytra is purplish to black and it is abundantly clothed with ferrugineus hairs. The head and thorax are finely to moderately coarsely, densely punctate, with many punctures confluent. The

elytral surface is densely, coarsely punctured, with the punctures not arranged in rows. The

basal-most antennomere is usually paler than the other segments. Adults are 7 to 15 mm long. It

may be mistaken for a large Enoclerus specimen, however the generic characters readily separate

them.

Biology: Bøving and Champlain (1920: 635) noted that T. apivorus is a pollen feeder and

has been recorded from the nests of bees.

It has been collected in Florida in insect flight traps with and without carbon dioxide baits,

Malaise traps, Steiner traps and McPhail traps. Adults have been collected from March through

November. It has been collected on Cirsium sp., Ambrosia artemisiifolia Linnaeus, Erigeron

quercifolius Poiret, Pterocaulon pycnostachyum (Michaux) Elliott, Bidens pilosa Linnaeus

(Asteraceae), Rhus copallinum Linnaeus (Anacardiaceae), Pinus elliottii Engelmann (Pinaceae),

Opuntia blooms (Cactaceae), Citrullus lunatus (Thunberg) Matsumura and Nakai

(Cucurbitaceae), Ilex glabra (Linnaeus) Gray (Aquifoliaceae), Melilotus albus Medikus, Cassia

sp. (Fabaceae), Polygonum hydropiperoides Michaux (Polygonaceae), Serenoa repens (Bartram)

Small, Sabal etonia Swingle (Arecaceae), Citrus sp. (Rutaceae), and Eriocaulon sp.

(Eriocaulaceae).

Distribution: Alabama, Colorado, District of Columbia, Florida, Georgia, Illinois,

Massachusetts, Mississippi, Nebraska, New Hampshire, New Jersey, New Mexico, New York,

North Carolina, Pennsylvania, South Carolina, Tennessee, Texas, Virginia and Canada (Wolcott,

1909: 98; Bøving and Champlain, 1920: 635; Wolcott, 1947: 82; Papp, 1960: 85; Barr, 1975:

11). Wickham and Wolcott (1912: 59) recorded it from Mount Tom, Massachusetts, which is

within 20 miles of Connecticut, and Webster, New Hampshire, which is within 40 miles of

Maine. Knull (1951: 301) stated that it should occur in Ohio. Foster (1976c: 26) reported

records for Louisiana and Texas, but believed them to be erroneous.

In Florida T. apivorus has been recorded from Alachua, Citrus, Clay, Duval, Franklin,

Hardee, Highlands, Indian River, Levy, Liberty, Marion, Miami-Dade, Okaloosa, Orange, Palm

Beach, Seminole, Taylor, and Walton Counties. Wickham and Wolcott (1912: 59) recorded it

from Crescent City, Putnam County, Florida.

Specimens examined (131): FLORIDA: Alachua County: Gainesville, 6-V-1922, 8-V-

1922, 22-V-1924, 25-V-1924, 28-V-1928, G.B. Merrill, 21-III-1947, 29-X-1946, H.V. Weems,

Jr., 19-VII-1991, M.C. Thomas, 10-XI-1946, [anon.], on Bidens pilosa, 24-VIII-1957, H.V.

Weems, Jr., on Rhus copallinum, 26-VIII-1953, H.V. Weems, Jr. (16, FSCA); Gainesville

Airport, 19-VII-1991, L. Heyer (1, FSCA); Gainesville, Austin Carey Forest, CO2-baited insect flight trap, 3-VI-1976, 12-VII-1976, G.B. Fairchild (2, FSCA); Highway 24 at Levy County line,

1-V-1978, R. Turnbow (1, RHTC); Monteocha, insect flight trap, 24-V-1977, 27-VII-1977, J.F.

Butler (2, FSCA); Newberry, 7-VIII-1959, R.E. Woodruff (1, FSCA); Windsor, on Pinus

elliottii, 10-VII-1986, T.W. Phillips (1, FSCA); 24-VI-1958, H.A. Denmark, 7-VIII-1955, H.V.

Weems, Jr., V-1970, L.A. Hetrick, on Melilotus alba, 24-V-1955, H.V. Weems, Jr. (5, FSCA);

Baker County: 1.3 miles north of Duval County along Route 90, 17-V-1992, L.R. Davis, Jr. (1,

FSCA); Bradford County: on Opuntia, 1-V-1959, H.V. Weems, Jr. (1, FSCA); Broward County:

1-VI-1958, D.R. Paulson (1, FSCA); Citrus County: Lecanto, 9-V-1983, P.M. Choate (1, FSCA);

Withlacoochee State Forest, 4-V-1988, C. Stevens (1, FSCA); Clay County: Camp Crystal, on

Pterocaulon undulatum, 6-V-1967, H.V. Weems, Jr. (1, FSCA); 2-VII-1960, H.A. Denmark (1,

FSCA); Collier County: Corkscrew Swamp, on Cirsium, 9-IV-1958, H.V. Weems, Jr. (1, FSCA);

Dixie County: 4 miles north of Old Town, 12-V-1979, 20-V-1978, E. Giesbert (2, FSCA); Duval

County: Jacksonville, McPhail trap, 7-VI-1950, 22-VI-1950, G.G. Norman (2, FSCA); Franklin

County: Apalachicola National Forest, Road 165, 19-V-1985, L.R. Davis, Jr. (1, FSCA);

Gilchrist County: Bell, on Citrulius vulgaris, 5-V-1954, H.M. Van Pelt (1, FSCA); Glades

County: 7 miles north of LaBelle, on Ambrosia artemisiifolia, 28-VI-1989, L. Maynard (1,

FSCA); Hardee County: Lily, in Steiner trap, 16-V-1969, R.H. Rhodes (1, FSCA); Highlands

County: Archbold Biological Research Station, 12-IV-1976, 13-IV-1976, W. Rosenberg, 24-IV-

1978, L.L. Lampert, Jr., Lake Placid, 22-IV-1983, H.L. Dozier (5, FSCA); Avon Park, 4-IV-

1953, K.V. Krombein (1, FSCA); Highlands Hammock State Park, 27-III-1950, H.V. Weems,

Jr., on Ilex glabra, 5-IV-1959, H.V. Weems, Jr., 14-IV-1990, M.C. Thomas (6, FSCA); near

Venus, 20-IV-1958, D.R. Paulson (1, FSCA); Venus, on Serenoa repens, 4-V-1961, R.E.

Woodruff, on Erigeron quercifolius, 21-III-1963, 27-III-1963, H.V. Weems, Jr. (5, FSCA);

Hillsborough County: Pebble Creek Golf Course, 24-V-1978 (3, FSCA); Indian River County:

Vero Beach, 4-VII-1932, 20-VI-1932, 30-VI-1932 E.M. Becton (3, FSCA); Levy County: 1 mile

east of Bronson, on Cassis, 22-VII-1992, C. Porter and L.A. Stange (1, FSCA); 13 miles

southwest of Williston, on Highway 121, 26-IV-1981, H.D. Baggett (1, FSCA); 3.8 miles southwest of Archer, 4-V-1988, P.E. Skelley, malaise trap in rosemary and turkey oak sandhill,

21-V-1988, P.E. Skelley (2, FSCA); Cedar Key, 15-VI-1980, L.A. Wood (1, FSCA); SR 24 at

Alachua County line, 1-V-1978, M.C. Thomas (1, FSCA); 19-VII-1958, H.V. Weems, Jr., on

Polygonum hydropiperoidea, 10-IX-1955, H.V. Weems, Jr. (2, FSCA); Liberty County: 20-VIII-

1944 (1, FSCA); Marion County: Ocala National Forest, NFR 75, Lake Delancy, on Pterocaulon pycnostachyum (Michaux), 5-VIII-1990, 11-VIII-1990, H.D. Baggett (2, FSCA); Village of

Rainbow Springs, 4-VII-1982, M.C. Thomas (1, FSCA); Miami-Dade County: Everglades

National Park, 10-IV-1955, F.W. Mead, 26-III-1976, L.L. Lampert, Jr., Long Pine Key, 29-IV-

1976, F.T. Hovore, Pinelands, 5-V-1978, 17-V-1978, R. Turnbow (4, FSCA; 4, RHTC);

Homestead, 3-IV-1962, R.E. Woodruff, on Sabal etonia, 27-IV-1933, O.D. Link (2, FSCA);

Kendall, SW 128th Street, 0.5 miles east of SW 137th Avenue, 26-VII-1990 (1, FSCA); Miami, on Citrus, 3-V-1932, O.D. Link (1, FSCA); 20-IV-1949, 22-IV-1952, 1950, [no date] (4, FSCA);

Okaloosa County: 3.2 miles southeast of Niceville, 27-V-1985, R. Turnbow (1, RHTC); 4.5 miles northwest of Holt, Florida A and M Research Station, Blackwater River State Forest, 31-

VIII-1978, L.A. Stange (1, FSCA); Orange County: FTU campus, Orlando, 9-VII-1979, 25-VII-

1972, 14-VI-1972, S.M. Fullerton (3, UCFC); SR 520, 5 miles east of SR 50, 9-VI-2006, (1,

UCFC); Wekiwa Springs State Park, burn zone 43/31, S22 T20S R28E, longleaf pine-turkey oak, Malaise Trap, 9-VI-2001, P. Russell and S.M. Fullerton (1, UCFC); Wekiwa Springs State

Park, burn zone WS-14a, S26 T20S R28E, longleaf pine-turkey oak, Black Pan Traps one foot above ground, 14-V-2001, P. Russell and S.M. Fullerton, Flor. Yel. Pan Traps one foot above ground, 8-VII-2001, 3-IV-2001, 14-IV-2001, P. Russell and S.M. Fullerton, Yellow Pan Traps one foot above ground, 10-VI-2001, P. Russell and S.M. Fullerton (5, UCFC); 15-V-1971, W.E.

Sterling, on Eriocaulon, 30-VI-1948, O.D. Link (2, UCFC; 1, FSCA); Clarcona, 17-VI-1932 (1,

FSCA); Osceola County: Kissimmee, 16-VI-1920, Reese (1, FSCA); Palm Beach County:

Jupiter, 16-VI-1966, B.K. Dozier (1, FSCA); Sarasota County: Myakka River State Park, Bee

Island, on log of Pinus, 30-IV-2002, J.L. Foltz (1, JMLC); Seminole County: lower Wekiwa

River State Preserve, burn zone LW-10, S39 T19S R29E, longleaf pine-turkey oak scrub, Flor.

Yel. pan traps one foot above ground, 14-IV-2001, P. Russell and S.M. Fullerton, red pan traps one foot above ground, 14-V-2001, P. Russell and S.M. Fullerton, white pan traps one foot above ground, 14-IV-2001, P. Russell and S.M. Fullerton (3, UCFC); Oviedo, 30-IV-1959, P.E.

Erierson (1, FSCA); on Opuntia blooms, 3-V-1960, C.O. Youtsey (1, FSCA); Taylor County:

Blue Springs Lake, 4-VI-1972, 5-VI-1972 R. Turnbow (5, FSCA); Union County: Santa Fe

River at Highway 241, 5-VII-1986, C.W. Mills, III (1, FSCA); Walton County: Eglin Air Force

Base, range 52A, 18-IV-1967, P.A. Thomas (1, FSCA); [county unknown]: Florida Agricultural

Exp. Station (4, FSCA).

Selected references: Germar, 1824: 81; Sturm, 1826: 59; Say, 1835: 164; Klug, 1842: 332; Sturm, 1843: 83; Spinola, 1844: 307, pl. 30, fig. 4; LeConte, 1849: 18; Gemminger and von Harold, 1869: 1742; Horn, 1876b: 231; Chittenden, 1890: 154; Hamilton, 1895: 335; Wickham, 1895: 249; Lugger, 1899: 149; Lohde, 1900: 59; Wolcott, 1909: 98, 1910b: 375, pl. 6, fig. 30; Wickham and Wolcott, 1912: 59; Wolcott and Chapin, 1918: 108; Bøving and Champlain, 1920: 635; Forbes, 1922: fig. 48; Balduf, 1935: 108; Wade, 1935: 108; Wolcott, 1944b: 60, 1947: 82; Knull, 1951: 301; Papp, 1960: 85; Ekis, 1975: 42; Foster, 1976c: 23; Barr, 1975: 11; Peck and Thomas, 1998: 86.

Figure 2-1. Enoclerus angustus

Figure 2-2. Distribution map of Enoclerus angustus

Figure 2-3. Enoclerus ichneumoneus

Figure 2-4. Distribution map of Enoclerus ichneumoneus

Figure 2-5. Enoclerus lunatus

Figure 2-6. Distribution map of Enoclerus lunatus

Figure 2-7. Enoclerus nigrifrons

Figure 2-8. Distribution map of Enoclerus nigrifrons

Figure 2-9. Enoclerus nigripes

Figure 2-10. Distribution map of Enoclerus nigripes

Figure 2-11. Enoclerus rosmarus

Figure 2-12. Distribution map of Enoclerus rosmarus

Figure 2-13. Placopterus thoracicus

Figure 2-14. Distribution map of Placopterus thoracicus

Figure 2-15. Priocera castanea

Figure 2-16. Distribution map of Priocera castanea

Figure 2-17. Thanasimus dubius

Figure 2-18. Distribution map of Thanasimus dubius

Figure 2-19. Trichodes apivorus

Figure 2-20. Distribution map of Trichodes apivorus

CHAPTER 3 ENOPLIINAE

Key to the Florida Genera of Enopliinae

1. Pronotum pink to orange, with two longitudinal vittae; elytra dull ...... Chariessa

1’. Pronotum variable, but never with two longitudinal vittae; elytra dull or shining ...... 2

2 (1). Pronotum with elevated regions; pronotum mottled brown; lichen-mimicking color pattern with a broad, irregularly margined midelytral white fascia ...... Pelonium leucophaeum (Fig. 3-11)

2’. Pronotum without elevated regions, but possibly with depressions; pronotum variable; elytra with or without a broad midelytral white fascia ...... 3

3 (2). Pronotum lacking lateral projections...... Neorthopleura thoracica (Fig. 3-9)

3’. Pronotum with lateral projections ...... Cregya

Genus Chariessa Perty, 1830

= Philyra Castelnau, 1836 = Brachymorphus Chevrolat, 1835

Chariessa is represented by four widely distributed species from North America (Opitz,

2002: 278). Chariessa is distinguished by dull, soft elytra which occasionally have yellow lateral and sutural margins, emarginated eyes and anterior tibiae which are serrulate outside.

Chariessa should not be mistaken for Pelonium, whose elytra appear polished, “harder” and have

color patterns that differ significantly (mimicking lichen in the Florida species). Neorthopleura

shares similar body size (at times), elytra and antennal structure. However, N. thoracica has an

entirely reddish pronotum, no color on the elytral margins, and has an occasional faint pale

median elytral fascia. Monophylla shares similar body form and elytra as well, but has an

elongate, spatulate apical antennomere while Chariessa has three laterally elongate apical

antennomeres.

Selected references: Perty, 1830: 109; Chevrolat, 1835, Fas. 7, No. 150; Castelnau, 1836: 53; Spinola, 1844: 347; LeConte, 1849: 32; Chevrolat, 1876: 7; LeConte and Horn, 1883: 219; Wickham, 1895: 248; Wolcott, 1908: 70, 1910c: 858; Bøving and Champlain, 1920:

608; Bradley, 1930: 107; Bøving and Craighead, 1931a: 57; Wolcott, 1947: 85; Opitz, 2002: 278.

Key to the Florida Species of Chariessa

1. Pronotal vittae thin, arched outward at the middle, the space between the vittae at middle much wider than the width of a single vitta; only known from Key Largo, Monroe County ...... C. floridana (Fig. 3-1)

1’. Pronotal vittae thick, subparallel, the space between the vittae much narrower than the width of a single vitta; widely distributed, but not extending to the southernmost seven counties of Florida ...... C. pilosa (Fig. 3-3)

Chariessa floridana Schaeffer, 1917 (Figs. 3-1, 3-2)

Chariessa floridana is similar to C. pilosa, but in C. floridana the pronotum much more

dominated by the pinkish coloration and the two longitudinal black vittae narrower and bent

outward from the center but converging normally anteriorly and posteriorly. Despite the

extensive variation of C. pilosa, it should not be confused with C. floridana. Both specimens

examined possessed yellow lateral and sutural elytral margins.

Biology: Few specimens of this elusive, firefly-mimicking species have been collected

(Mawdsley, 1992: 28). In Florida adults have been collected in May and July.

Distribution: Schaeffer (1917: 133) and Wolcott (1947: 85) recorded it from Key Largo,

Florida (the type locality). Papp (1960: 86) and Barr (1975: 16) only recorded this species from

Florida. No literature has suggested this species to occur elsewhere.

Specimens examined (2): FLORIDA: Monroe County: upper Key Largo, 30-V-1976,

M.C. Thomas and J.H. Frank (1, FSCA); Key Largo, 17-VII-1959, W.W. Warner (1, FSCA).

Selected references: Schaeffer, 1917: 133; Wolcott, 1947: 85; Papp, 1960: 86; Barr, 1975: 16; Mawdsley, 1992: 28; Peck and Thomas, 1998: 86.

Chariessa pilosa Forster, 1771 (Figs. 3-3, 3-4)

= Chariessa pilosa var. marginata Say, 1823

Chariessa pilosa has a pinkish pronotum with two black vittae and may have yellow sutural and lateral elytral margins accentuating the otherwise black color. Its body and elytra share a soft, smooth appearance with no form of shininess. It shares the body form and similar antennae with Monophylla, Pelonium, and Neorthopleura (see the genus discussion above for distinguishing characters). Adults are 7 to 13 mm long. Bøving and Champlain (1920: 608) described the larvae and compared them to some related taxa.

Biology: Chariessa pilosa has been recorded from oak, hickory, walnut, pear, sassafras, linden, grape, birch, chestnut, elm, and coniferous trees infested with Saperda tridentata Olivier,

Xylotrechus colonus (Fabricius), Neoclytus acuminatus (Fabricius) (Coleoptera: Cerambycidae),

Chrysobothris femorata (Olivier) (Coleoptera: Buprestidae), Magdalis sp. (Coleoptera:

Curculionidae), and Scolytus multistriatus (Marsham) (Coleoptera: Curculionidae: Scolytinae)

(Hoffman, 1940: 56; Knull, 1951: 307). After infiltrating the galleries of its prey species it exploits the prey species pupal cell, as it does not construct one of its own (Bøving and

Champlain, 1920: 638).

Wickham and Wolcott (1912: 66) noted that it has been reared from elm. Gosling (1980:

74) observed it at ultraviolet lights in oak-hickory habitats. It is considered to be a mimic of fireflies (Mawdsley, 1992: 28).

Adults are diurnal and have been collected in Florida from March to July. Specimens have been recorded collected at mercury vapor lights and blacklights, from ethanol-baited Lindgren funnel traps and from flight-intercept or Malaise traps. One specimen was collected on

“persimmon stump sprouts”.

Distribution: Alabama, Arkansas, Connecticut, District of Columbia, Florida, Georgia,

Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts,

Michigan, Mississippi, Missouri, Nebraska, New Hampshire, New Jersey, New York, Ohio,

Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Utah, Wisconsin, and

Canada (Wolcott, 1909: 100, 1947: 85; Bøving and Champlain, 1920: 638; Knull, 1951: 307;

Papp, 1960: 85; Barr, 1975: 16; Gosling, 1980: 74). Wickham and Wolcott (1912: 66) recorded this species from Bayfield, Wisconsin, which is within 40 miles of Minnesota and 75 miles south of Ontario.

In Florida it has been recorded from Alachua, Bradford, Dixie, Gadsden, Gilchrist, Gulf,

Hernando, Highlands, Hillsborough, Indian River, Jefferson, Leon, Levy, Liberty, Marion,

Orange, Polk, Putnam, Santa Rosa, Seminole, Wakulla, and Walton Counties. Peck and Thomas

(1998: 86) included Monroe County in the distribution of C. pilosa. Examination of the specimen cited showed it to be the much rarer C. floridana. With this clarification, there are no records of C. pilosa in south Florida (i.e., the seven southernmost counties).

Specimens examined (132): FLORIDA: Alachua County: 9 miles northwest of

Gainesville, U.F. Horticultural Unit, malaise trap, 20-IV-1977, H.N. Greenbaum (1, FSCA);

Austin Cary Forest, insect flight trap, 21-IV-1986, G.B. Fairchild and Roberts, 21-III-1982, G.B.

Fairchild and H.V. Weems, Jr. (2, FSCA); Gainesville, VI-2003, J.M. Leavengood, Jr., 21-V-

1978, M.C. Thomas (1, JMLC; 1, FSCA); Payne's Prairie State Preserve, insect flight trap, 16-V-

1984, G.B. Fairchild and H.V. Weems, Jr. (1, FSCA); Bradford County: SR 100, 8km west of

Starke, 30-VI-1980, A. Wilkening (1, FSCA); Dixie County: 3.5 miles north of Old Town, 27-

IV-1979, 8-V-1978, 9-V-1978, M.C. Thomas (4, FSCA); 4 miles north of Old Town, 20-V-1978,

E. Giesbert, 7-V-1979, R. Turnbow (19, FSCA; 1, RHTC); 5 miles north of Old Town, on persimmon stump sprouts, 23-V-1978, R. Woodruff and P.M. Choate, Jr. (1, FSCA); highway

349, 3 miles south of junction, 351, 7-V-1978, R. Turnbow (8, RHTC); Old Town, 20-V-1978,

G.B. Edwards (2, FSCA); Gadsden County: Aspalaga Landing, Aspalaga Rd., 9-V-2005, E.

Nearns (1, FSCA); Gilchrist County: vic. Heart Springs, 30-V-1988, R. Turnbow (1, RHTC);

Gulf County: 15.1 miles northeast of Port St. Joe, 29-V-1989, R. Turnbow (1, RHTC); Hernando

County: Withlacoochee State Forest, Richloam Tract, 8-IV-1994, 9-IV-1992, M.C. Thomas (3,

FSCA); Highlands County: Archbold Biological Station, insect flight trap, 4-VI-1978, H.V.

Weems, Jr. and L. Klein, malaise trap, 25-IV-1980, L.L. Lampert, Jr. (2, FSCA); Hillsborough

County: USF Campus (1, FSCA); Jefferson County: junction of highways 59 and 98, 30-V-1988,

4-VI-1988, R. Turnbow (2, RHTC); Monticello, 17-IV-1994, 24-IV-1994, 1-V-1994, 8-V-1994,

21-V-1994, 28-V-1994, Buck B. (8, FSCA); Leon County: Tallahassee, 4-VII-2002, G.J.

Wibmer, at ultraviolet light trap, 29-VI-2002, G.J. Wibmer and C.W. O'Brien (3, FSCA); Levy

County: 3.8 miles southwest of Archer, malaise trap in rosemary and turkey oak sandhill, 10-IV-

1988, 17-IV-1988, 23-IV-1988, P.E. Skelley (3, FSCA); Liberty County: Apalachicola Bluffs and Ravines Preserve, 4-V-2003, R. Turnbow (2, RHTC); Highway 12, 17 miles south of Bristol,

29-V-1989, R. Turnbow (1, RHTC); Torreya State Park, ethanol-baited Lindgren funnel trap, V-

2007, VI-2007, J.M. Leavengood, Jr. (4, JMLC); Torreya State Park, 6-V-1979, 7-V-1989, M.C.

Thomas, 16-V-1971, H.V. Weems, Jr. and G.B. Fairchild, flood plain forest, 5-IV-2003, E.

Nearns (3, FSCA; 1, JMLC); Marion County: 8 miles southeast of Interlachen, 21-III-1953, B.K.

Dozier (1, FSCA); junction of highways 40 and 314A, 17-V-1986, R. Turnbow (2, RHTC);

Route 316, Oklahawa River, under bridge, at light, 5-V-2002, M. Hrabovsky (1, FSCA); Orange

County: Rk Spr Rn St Res, Spine/Oak Scrub, Malaise Trap, 3-V-1995, 24-V-1995, S.M.

Fullerton (2, UCFC); University of Central Florida, Orlando, Cypress Forest, Malaise Trap, 29-

III-1999, 12-IV-1999, 17-V-1999, 2-VI-1999, 8-VI-1999, 14-VI-1999, 21-VI-1999, P. Russell and S.M. Fullerton (8, UCFC); University of Central Florida, Orlando, LLP-Sand Pine, Turkey

Oak, Malaise Trap, 17-V-1996, 30-V-1997, 11-VI-1997, 18-VI-1997, 10-VII-1997, S.M.

Fullerton (5, UCFC); University of Central Florida, Orlando, Longleaf Pine-Saw Palmetto,

Malaise Trap, 11-VI-1997, S.M. Fullerton (1, UCFC); University of Central Florida, Orlando,

Maidencane Marsh, Malaise Trap, 2-VI-1999, 14-VI-1999, P. Russell and S.M. Fullerton (4,

UCFC); University of Central Florida, Orlando, Pond Pine Community, Dahoon Holly, Malaise

Trap, 17-V-1999, 24-V-1999, 2-VI-1999, 8-VI-1999, 7-VII-1999, 13-VII-1999, 19-VII-1999, P.

Russell and S.M. Fullerton (9, UCFC); University of Central Florida, Orlando, Sand Pine-

Rosemary Scrub, Malaise Trap, 13-VI-1993, S.M. Fullerton (1, UCFC); Walt Disney World, C-4

Stout Site, S 15, 16 T 24S R 27E, xeric oak/flatwoods, Malaise Trap, 12-19-V-1998, Z. Prusak and S.M. Fullerton (1, UCFC); Walt Disney World, MW - 7 (unburned), S 22 T 24S R 27E, sand pine/oak scrub, Malaise Trap, 29-V-5-VI-1997, Z. Prusak and S.M. Fullerton (1, UCFC); Polk

County: Lake Marion Estates, 1-V-1999, R. Turnbow, 1-V-1999, M.C. Thomas (1, RHTC; 1,

FSCA); Tiger Creek Preserve, Malaise trap, 19-IV-2007, D.T. Almquist (1, JMLC); Putnam

County: 2-V-1959, in spider web 11-V-1959, H.V. Weems, Jr. (2, FSCA); Santa Rosa County:

11.7 miles east of Gulf Breeze, 18-VI-1988, R. Turnbow (1, RHTC); 4 miles north of Munson,

28-IV-1984, R. Turnbow (2, RHTC); Gulf Breeze, ethanol-baited Lindgren funnel trap, 4-V-

2007, 20-VI-2007, C. Street (3, CAPS); Seminole County: Econ. Wild. Area, scrub oak/saw palmetto burn site, Malaise Trap, 6-V-2000, 15-VII-2000, 26-VIII-2000, T. Smith, P. Russell and S.M. Fullerton (3, UCFC); Oviedo, rural yard, Malaise Trap, 5-VI-1994, S.M. Fullerton (1,

UCFC); Wakulla County: 6 miles east of Newport, 4-VI-1988, R. Turnbow (2, RHTC); Walton

County: highway 285 at Shoal River, 5-V-1985, R. Turnbow (1, RHTC).

Selected references: Forster, 1771: 49; Say, 1823: 187; Klug, 1842: 360; Spinola, 1844: 356, 358, pls. 34, 36, figs. 4, 5; LeConte, 1849: 32; Drury, 1879: 171; Chittenden, 1890: 155; Hamilton, 1895: 335; Wickham, 1895: 252; Lohde, 1900: 104; Felt, 1906: 504; Wolcott, 1908: 72; Wolcott, 1909: 100, 1910c: 859; Wickham and Wolcott, 1912: 66;

Bøving and Champlain, 1920: 608, 638; Blackman and Stage, 1924: 46; Hoffmann, 1940: 56, 1942: 7; Beal and Massey, 1945: 69, 92; Wolcott, 1947: 85; Craighead, 1950: 199; Knull, 1951: 306, pl. 10, fig. 38; Papp, 1960: 85; Ekis, 1975: 52; Barr, 1975: 16; Gosling, 1980: 74; Mawdsley, 1992: 28; Peck and Thomas, 1998: 86; Opitz, 2002: figs. 80, 88, 89, 119.

Genus Cregya LeConte, 1861

Cregya is represented by three described species in the eastern, central and southern United

States to Texas (Barr 1962: 127). Two described species are known from Florida. It is distinguished by its emarginated eyes, 11-segmented antennae and its conspicuously expanded, angulate pronotal sides. Cregya is the only genus of the subfamily with eleven antennomeres.

Bøving and Champlain (1920: 606) described the larvae and compared them to some related taxa.

Selected references: LeConte, 1861: 197; Wolcott, 1910b: 383, 1910c: 860; Gahan, 1910: 75; Bøving and Champlain, 1920: 606; Bøving and Craighead, 1931a: 57; Wolcott, 1947: 86; Barr, 1962: 127; Opitz, 2002: 279.

Key to the Florida Species of Cregya

1. Elytra black, with sutural and lateral margins pale; widely distributed ...... C. oculata (Fig. 3-5)

1’. Elytra pale with black maculations; only known from Palm Beach County in Florida ...... C. quadrisignata (Fig. 3-7)

Cregya oculata (Say), 1835 (Figs. 3-5, 3-6)

= Pelonium marginipenne Spinola, 1844 = Enoplium circumcinctum Spinola, 1844 = Pelonium lineolatum Gorham, 1883

The elytral markings of C. oculata are rather consistent with the exception of variably

reduced pale elytral margins. When reduced, these pale borders may only reach halfway from

the apex toward the base of each elytron (or anywhere between the halfway point and the base).

The mouth parts, basal antennomeres, prothorax, and parts of the legs are yellow. The pronotum

varies considerably from being entirely piceous to having variously shaped maculations. The

body is moderately clothed with erect and semierect hairs, longest on head and thorax. Adults are 4.0 to 7.0 mm long. Despite the extreme variation in pronotal maculations, C. oculata is rather easily diagnosed from C. quadrinotata by the color of the elytra. Bøving and Champlain

(1920: 606) described the larvae and compared them to some related taxa.

Biology: Cregya oculata has been recorded from pitch pine, sumac, chestnut, Virginia creeper infested with Pogonocherus sp., Leiopus sp., Euderces picipes (Fabricius), Ecyrus dasycerus (Say) (Cerambycidae), and Pityophthorus sp. (Coleoptera: Curculionidae: Scolytinae)

(Knull, 1951: 309; Bøving and Champlain, 1920: 639). Gosling (1980: 74) observed it at ultraviolet lights in oak-hickory habitats.

Adults have been collected in Florida from April through July and September. Specimens have been collected from mercury vapor lights, blacklights, Steiner traps and Malaise traps.

Distribution: Alabama, District of Columbia, Florida, Georgia, Indiana, Illinois, Kansas,

Kentucky, Louisiana, Massachusetts, Michigan, New Jersey, New York, North Carolina,

Pennsylvania, Ohio, South Carolina, Texas, Virginia, and West Virginia (Wolcott, 1909: 101,

1910b: 390, 1947: 86; Wickham and Wolcott, 1912: 67; Bøving and Champlain, 1920: 639;

Knull, 1951: 308; Papp, 1960: 86; Barr, 1975: 17; Gosling, 1980: 74).

In Florida it has been recorded from Alachua, Bay, Collier, Dixie, Gulf, Hardee, Hernando,

Highlands, Indian River, Jefferson, Levy, Liberty, Marion, Monroe, Okaloosa, Orange, Santa

Rosa, Seminole, Volusia, and Wakulla Counties. Wolcott (1910b: 390) recorded it from Tampa,

Hillsborough County, Florida.

Specimens examined (39): FLORIDA: Alachua County: Gainesville, at light, 30-V-1947,

1-VI-1959, 11-VI-1948, H.V. Weems, Jr. (3, FSCA); SE Kincaid Rd, edge old field, xeric oak hammock, 6 m Malaise trap, 8-15-V-1999, 13-26-VI-1999, B.D. Sutton (2, FSCA); SE Kincaid

Rd, 1 mile north of Payne's Prairie, edge old field/xeric oak hammock, 6 m Malaise trap, 24-IV-

4-V-1999, 24-IV-4-V-1998, 4-11-V-1998, 17-24-V-1998, 24-31-V-1998, B.D. Sutton (5,

FSCA); 6-V-1969, M.L. May (1, FSCA); Bay County: 2 miles north of West Bay, 10-V-1987,

16-V-1987, R. Turnbow (3, RHTC); 5.4 miles north of Lynn Haven, 29-V-1989, R. Turnbow (1,

RHTC); Collier County: Highway 41, east of 29, 18-IV-1998, R. Morris (1, FSCA); Dixie

County: 4 miles north of Old Town, 20-V-1978, E. Giesbert (1, FSCA); Gulf County: 10 miles west of Wewahitchka, 7-V-1988, R. Turnbow (1, RHTC); Hardee County: Ona, Steiner trap, 14-

VII-1967, R.H. Rhodes (1, FSCA); Hernando County: 1 mile north of the junction of highways

50 and 301, 27-IV-1991, R. Turnbow (1, RHTC); Withlacoochee State Forest, Richloam Tract,

9-IV-1992, M.C. Thomas, 8-IV-1994, 27-IV-1991, R. Turnbow (1, FSCA; 2, RHTC); Highlands

County: 2 miles south of Sebring, 10-IV-1990, R. Turnbow (1, RHTC); Jefferson County: junction of highways 59 and 98, 30-V-1988, 4-VI-1988, R. Turnbow (2, RHTC); Levy County: 5 miles north of Otter Creek, 13-V-1979, E. Giesbert (1, FSCA); Liberty County: Torreya State

Park, 7-V-1989, C.W. Mills, III, 25-VI-1989, R. Turnbow (1, FSCA; 1, RHTC); Monroe

County: Big Pine Key, 29-IV-1977, E. Giesbert (1, FSCA); Okaloosa County: Eglin Air Force

Base, Santa Rose Island, west of Destin, blacklight trap in dunes, 17-VI-1995, P.E. Skelley (1,

FSCA); Orange County: University of Central Florida, MacKay Tract, Orlando, Sawgrass

Marsh, red maple, Malaise Trap, 19-IV-1999, 17-V-1999, 24-V-1999, P. Russell and S.M.

Fullerton (3, UCFC); Santa Rosa County: 11.7 miles east of Gulf Breeze, 18-VI-1988, R.

Turnbow (1, RHTC); Seminole County: Econ. Wild. Area, scrub oak/saw palmetto burn site,

Malaise Trap, 10-IX-2000, T. Smith, P. Russell and S.M. Fullerton (1, UCFC); Volusia County:

Oak Hill, Steiner trap, 8-IV-1963, R.H. Rhodes (1, FSCA); Wakulla County: 6 miles east of

Newport, 4-VI-1988, 11-VI-1988, R. Turnbow (2, RHTC).

Selected references: Say, 1835: 163; Klug, 1842: 387; Spinola, 1844: 363, pl. 35, fig. 6; LeConte, 1849: 32; Melsheimer, 1853: 84; LeConte, 1859b: 639; LeConte, 1865: 98; Drury, 1879: 171; Gorham, 1883: 191, pl. 9, fig. 14; Henshaw, 1885: 82; Hamilton, 1895: 335; Lohde, 1900: 103; Schenkling, 1906: 317; Wolcott, 1909: 101, 1910b: 389, 1910c: 860; Wickham and Wolcott, 1912: 67; Bøving and Champlain, 1920: 606, 639; Knull, 1932: 43; Wolcott, 1947: 86; Craighead, 1950: 203; Knull, 1951: 309, pl. 11, fig. 45; Papp, 1960: 86; Ekis, 1975: 53; Barr, 1975: 17; Gosling, 1980: 74; Peck and Thomas, 1998: 86; Opitz, 2002: 279, figs. 83, 97.

Cregya quadrinotata Spinola, 1844 (Figs. 3-7, 3-8)

= Pelonium quadrisignatum Spinola, 1844 (?) = Pelonium bilineicolle Chevrolat, 1874 (?)

While the maculations in C. quadrinotata are variable, the head is typically black. The mouth parts, a frontal maculation and the inner margin of the eyes tend to be pale yellow. The abdominal and thoracic sternites, tarsi, a large humeral maculation and a rounded maculation at the apical fourth of the elytra are usually black. Its body form is otherwise similar to C. oculata.

Adult are 4.0 to 5.5 mm long.

There is some uncertainty whether C. quadrinotata and C. quadrisignata are two separate species or not. Papp (1960: 86) referred to C. quadrisignata quadrinotata—but did not cite where this assignment was originally posited.

Biology: Bøving and Champlain (1920: 639) recorded it from dead black walnut branches.

Distribution: Wolcott (1909: 102, 1910b: 393, 1927: 3, 1947: 86) recorded C. quadrisignata from Texas, Mexico, Guatemala, Honduras, Nicaragua, Panama and Columbia.

Barr (1975: 17) recorded C. quadrinotata from Texas, and recorded C. quadrisignata from

Panama, Guatemala and Honduras. Papp (1960: 86) recorded C. quadrisignata quadrinotata from Texas. Bøving and Champlain (1920: 639) recorded C. quadrinotata it from West

Virginia. It has been collected in Palm Beach County in Florida.

Specimens examined: No Florida specimens were available for observation.

Selected references: Spinola, 1844: 365, pl. 33, fig. 3; LeConte, 1849: 31; Haldemann, 1853: 362; Chevrolat, 1874: 327; Gorham, 1883: 190; Lohde, 1900: 105, 107; Schenkling, 1903a: 107; Wolcott, 1909: 102, 1910b: 392; Bøving and Champlain, 1920: 639; Wolcott, 1927: 3, 1947: 86; Papp, 1960: 86; Ekis, 1975: 54; Barr, 1975: 17.

Genus Neorthopleura Barr, 1976

There are seventeen species of Neorthopleura in the Western Hemisphere, three of which represent the North American fauna across the eastern, central and western United States and

Mexico (Barr, 1962: 126). The genus is distinguished by its emarginated eyes, 11-segmented antennae and the broadly rounded sides of the pronotum, which is typically reddish. The dark, soft elytra may have a faint, pale transverse median fascia. Measuring from 3 to 12 mm in length, they vary greatly in size, the males often smaller than females.

This genus was originally named Dermestoides Schaeffer in 1771. However, the name was not binary and was discarded. They are rather active on infested trees in the evening. Opitz

(2002: 279) reported associations with numerous species of Cerambycidae (Coleoptera),

Curculionidae (Coleoptera) and Chrysobothris spp. (Coleoptera: Buprestidae) in oak, chestnut, and other hardwoods.

Bøving and Champlain (1920: 617) described the larvae under the name Orthopleura and compared them to some related taxa.

Selected references: Schaeffer, 1771: pl. 220, fig. 4; Spinola, 1844: 80; LeConte and Horn, 1883: 219; Wickham, 1895: 248; Wolcott, 1910b: 393, 1910c: 858; Bradley, 1930: 107; Bøving and Champlain, 1920: 617; Bøving and Craighead, 1931a: 56; Wolcott, 1943: 134; Barr, 1962: 126, 1976b: 2; Opitz, 2002: 279.

Neorthopleura thoracica (Say), 1823 (Figs. 3-9, 3-10)

= Tillus damicornis Fabricius, 1798 = Enoplium punctatissimum Chevrolat, 1843 = Orthopleura nigripennis Spinola, 1844 = Orthopleura bimaculata Melsheimer, 1845 = Pelonium pennsylvanicum Chevrolat, 1874

The pronotum of this N. thoracica is often reddish, only margined with the black to bluish- black which covers the remainder of its body. But it can also be entirely black to bluish-black.

Likewise, the normally black to bluish-black (save the pronotum) body can instead assume a

testaceous color. Lastly, specimens of this species may have a more or less distinct post-median pale fascia on the elytra. Despite the numerous variations, it is rather simple to distinguish from the other genera.

The antennal club of males measures more than two-thirds of the total antennal length and

more than one-half the total antennal length in females. It has toothed tarsal claws and cylindrical apical maxillary and labial palpomeres. Neorthopleura thoracica exhibits much variation in size and color. In reference to its range in size, Wickham and Wolcott (1912: 67)

wrote that “this insect is the most variable in size of any beetle I know”. Adults measure 5 to 14

mm long. The generic characters should be used to distinguish Neorthopleura from similar

species in Florida.

Bøving and Champlain (1920: 618) described the larvae and compared them to some

related taxa.

Biology: This nocturnal species has been recorded from oak, hickory, grape, locust, elm,

persimmon, ash, sassafras, and chestnut infested with Chrysobothris femorata (Olivier)

(Coleoptera: Buprestidae), Obrium sp., Neoclytus confusus Van Dyke, Tilliclytus geminatus

(Haldemann), Phymatodes aereus (Newman) (Coleoptera: Cerambycidae), and Curculionidae

(Coleoptera) (Bøving and Champlain, 1920: 639; Knull, 1951: 305). Gosling (1980: 74)

observed it at ultraviolet lights in oak-hickory habitats.

Adults have been collected in Florida from February through October. Specimens have

been collected at blacklights, halogen lights, mercury vapor lights, ultraviolet lights, Lindgren

funnel traps baited with alpha-beta pinene, ethanol, both turpentine and frontalin, Malaise traps

and under bark. It has been reared from Lysiloma latisiliqua (Linnaeus) (Fabaceae) and

hackberry.

Distribution: Arkansas, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa,

Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New

Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Texas, Virginia, the

Bahamas, Cuba, Mexico, Ontario, Quebec, the West Indies (Wolcott, 1909: 101, 1910b: 396;

Bøving and Champlain, 1920: 639; Wolcott, 1947: 84; Knull, 1951: 305; Vaurie, 1952a: 4; Papp,

1960: 85; Barr, 1975: 15, 1976b: 8). Wickham and Wolcott (1912: 67) recorded it from Onaga,

Kansas, which is 30 miles south of Nebraska, and Fort Lee, New York, which is within 30 miles

of Connecticut. There are records of N. thoracica from California, but Barr (1950a: 513)

suspected that the inclusion of “Lower California” may have simply been due to Horn’s report of its collection in San Jose del Cabo, making the record erroneous since he observed no specimens from California.

In Florida it has been recorded from Alachua, Bay, Brevard, Broward, Clay, Collier, Dixie,

Duval, Escambia, Gadsden, Hernando, Highlands, Hillsborough, Indian River, Leon, Levy,

Liberty, Manatee, Marion, Miami-Dade, Monroe, Orange, Pinellas, Santa Rosa, Sarasota,

Seminole, Sumter, Taylor, Volusia, Wakulla, Walton, and Washington Counties.

Specimens examined (229): FLORIDA: Alachua County: 2 miles north of Cross Creek,

on Highway 325, 7-VII-1991, C.W. Mills, III (2, FSCA); 2 miles northwest of Gainesville,

blacklight, 20-IV-1974, J.B. Heppner (1, FSCA); 6 miles southwest of Gainesville, blacklight

trap, 13-III-1975, 25-III-1975, 8-IV-1975, 1-VI-1975, L.R. Davis, Jr., 21-IV-1968, R.E.

Woodruff, 13-VI-1956, H.A. Denmark (6, FSCA); Gainesville, VI-1968, 3-VI-1980, 8-VII-1980,

5-IX-1977, 5-IX-1986, 12-IX-1981, 17-IX-1981, 30-IX-1981, 2-X-1980, L.A. Hetrick, 15-IX-

1987, P. Landolt, 24-XI-1977, [anon.], 4-VII-1978, M.C. Thomas, VIII-2003, IX-2003, J.M.

Leavengood, Jr. (13, FSCA; 2, JMLC); Gainesville, Beville Hts., blacklight trap, 13-IV-1980, 2-

VII-1980, L.A. Stange (2, FSCA); Gainesville, Devil's Millhopper State Park, Millhopper Drive,

Turpentine/frontalin-baited Lindgren funnel trap, IV-2005, J.M. Leavengood, Jr. (1, JMLC);

Gainesville, Doyle Conner building, at light, 14-IV-1990, 1-VI-1989, 6-VIII-1990, P.E. Skelley, blacklight trap, 24-VIII-1972, F.W. Mead (4, FSCA); Gainesville, Hogtown Creek and I-75, light trap, 19-VI-1988, P.E. Skelley (1, FSCA); Gainesville, NW 42nd Terr. and 34th Pl., ethanol-baited Lindgren funnel trap, 1-VI-2006, J.L. Foltz and J.M. Leavengood, Jr. (1, JMLC);

Gainesville, SE Kincaid Rd, 1 mile north of Payne's Prairie, old field dry oak hammock, Malaise trap, 14-22-VI-1998, B.D. Sutton (1, FSCA); Gainesville, University of Florida, Surge/Natural

Area Drive, Turpentine/frontalin-baited Lindgren funnel trap, IV-1998, J.L. Foltz (1, JMLC);

NW Gainesville, at light, 28-IV-2002, 11-VII-2002, E. Nearns, reared from hackberry, 26-V-

2002, E. Nearns (3, JMLC); Payne's Prairie State Preserve, gate 9B on Highway 234, malaise trap, 26-IV-1996, G.E. Steck and B.D. Sutton (1, FSCA); 19-IV-1955, R.A. Morse (1, FSCA);

Bay County: Panama City, ethanol-baited Lindgren funnel trap, 14-IX-2006, 3-X-2006, L. Smith

(1, JMLC; 1, CAPS); Brevard County: Titusville, SR 405, Enchanted Forest Sanctuary, White

Trail, xeric oak hammock, Malaise Trap, 21-III-4-IV-2001, 13-27-VI-2001, 11-25-VII-2001, P.

Russell, Z. Prusak and S.M. Fullerton (3, UCFC); Broward County: ethanol-baited Lindgren funnel trap, 29-V-2007, M. Meadows (2, CAPS); Clay County: Goldhead Branch State Park, 25-

VIII-1994, C. Porter and L.A. Stange (1, FSCA); Collier County: Collier-Seminole State Park, at mercury vapor and black light, 15-IV-1989, R. Turnbow, M.C. Thomas (1, RHTC; 1, FSCA);

Dixie County: 5 miles north of Old Town, 12-V-1979, 20-V-1978, E. Giesbert (7, FSCA); Duval

County: Jacksonville, ethanol-baited Lindgren funnel trap, 18-V-2007, 7-VI-2007, A. Silagyi and A. Johnson, A. Johnson (2, CAPS); Escambia County: ethanol-baited Lindgren funnel trap,

20-VI-2007, C. Street (2, CAPS); Gadsden County: Aspalaga Landing, at blacklight, 5-V-2007,

J.M. Leavengood, Jr. (2, JMLC); Hernando County: Withlacoochee State Forest, Goat Road, 22-

V-1992, 14-VI-1991, R. Morris, at blacklight, 4-VI-1991, R. Morris and J. Heuther (3, FSCA);

Withlacoochee State Forest, Richloam Tract, 9-IV-1992, M.C. Thomas, 18-IV-1992, 20-IV-

1992, R. Turnbow (4, FSCA); Highlands County: Archbold Biological Station, at mercury vapor and black light, 9-IX-1999, 10-IX-1999, R. Turnbow (4, RHTC); Archbold Biological Station, at ultraviolet light, 20-IV-1979, 26-IV-1976, L.L. Lampert, Jr. (3, FSCA); Archbold Biological

Station, malaise trap, 9-IV-1980, L.L. Lampert, Jr. (1, FSCA); Archbold Biological Station, 1-V-

1978, F. Hovore, 17-IV-1981, N.M. Downie, 26-IV-1978, 27-IV-1978, 28-IX-1977, 30-IX-1977,

L.L. Lampert, Jr. (6, FSCA); Archbold Biological Station, 10 miles south of Lake Placid, 2-V-

1975, J.B. Heppner (1, FSCA); Archbold Biological Station, 8 miles south of Lake Placid, blacklight trap, 26-V-1997, P.E. Skelley (2, FSCA); Hillsborough County: Lithia, ethanol-baited

Lindgren funnel trap, VI-2007, J.M. Leavengood, Jr. (1, JMLC); Tampa, ethanol-baited

Lindgren funnel trap, 3-V-2007, 1-VI-2007, 3-VII-2007, 21-IX-2006, 11-X-2006, D. Gaskill, 20-

VI-2007, Patterson, III-2007, J.M. Leavengood, Jr. (6, CAPS; 4, JMLC); Indian River County:

Vero Beach, 12-V-1964, 16-V-1964, W.L. Bidlingmayer (2, FSCA); [County unknown], Royal

Palm Park, V-1948 (1, FSCA); Leon County: Tall Timbers Research Station, 30-IX-1989, P.E.

Skelley (1, FSCA); Tallahassee, ethanol-baited Lindgren funnel trap, 4-V-2007, M. Bentley (1,

CAPS); Tallahassee, near I-90 and 319, at blacklight, 22-VI-1987, P.E. Skelley (1, FSCA);

Tallahassee, UV light trap, 12-14-VIII-2002, M. Haseeb, 28-VII-2002, 23-VIII-2002, G.J.

Wibmer, 6-VII-2002, C.W. O'Brien and G.J. Wibmer, 7-IX-2002, C.W. O'Brien (11, FSCA);

Levy County: 3.8 miles southwest of Archer, blacklight trap, 20-V-1988, P.E. Skelley (1,

FSCA); Liberty County: 14.8 miles north of Sumatra, 7-V-1988, R. Turnbow (2, RHTC);

Apalachicola National Forest, ethanol-baited Lindgren funnel trap, V-2007, J.M. Leavengood, Jr.

(1, JMLC); Torreya State Park, at light, 15-VII-1987, 17-VII-1987, Matthews and P.E. Skelley,

at mercury vapor and black light, 6-V-1989, R. Turnbow (3, FSCA; 1, RHTC); Torreya State

Park, ethanol-baited Lindgren funnel trap, V-2007, VI-2007, J.M. Leavengood, Jr. (3, JMLC);

Torreya State Park, 10-V-2005, E. Nearns, 16-V-1964, R.E. White, 6-V-1989, R. Turnbow, 4-6-

V-2007, J.M. Leavengood, Jr. (2, FSCA; 1, RHTC; 3, JMLC); Manatee County: Oneco, P.

Dillman (1, FSCA); Marion County: Lake Eaton, blacklight trap, 8-IV-1975, P. Drummond (1,

FSCA); McIntosh, ethanol-baited Lindgren funnel trap, IV-2007, J.M. Leavengood, Jr. (4,

JMLC); Ocala, blacklight trap, 25-VIII-1977, M.C. Thomas (1, FSCA); Miami-Dade County:

Camp Mahatchee, Matheson Hammock, 9-IV-1994, R. Turnbow (1, FSCA); Everglades

National Park, Long Pine Key, at halogen light, 16-V-1991, R. Morris (1, FSCA); Matheson

Hammock, 26-V-1962, R.E. Woodruff (1, FSCA); Miami, alpha-beta pinene baited Lindgren

funnel trap, 31-V-2007, M. Meadows (1, CAPS); Miami, blacklight trap, 15-IV-1960, 22-VII-

1960, P.E. Briggs (2, FSCA); Miami, ethanol-baited Lindgren funnel trap, 2-IV-2007, 31-V-

2007, M. Meadows, IV-2007, V-2007, VI-2007, J.M. Leavengood, Jr. (2, CAPS; 4, JMLC); V-

1962 (1, FSCA); Monroe County: J. Pennekamp Coral Reef State Park, north Key Largo, telephone pole #106, 15 watt blacklight, 29-V-1997, P.E. Skelley (1, FSCA); Key Largo, at mercury vapor and black light, 13-V-1986, R. Turnbow (1, RHTC); Key Largo, emerged 25-III-

1978, emerged 15-V-1978, R. Turnbow, emerged 9-VII-1961, B.K. Dozier (3, FSCA); Key

Largo, emerged from Lysiloma 12-III-1962, B.K. Dozier (1, FSCA); Key Largo, emerged from

Lysiloma latisiliqua (L.) 18-II-1978, 28-II-1978, 31-III-1978, 15-V-1978, R. Turnbow (1, FSCA;

5, RHTC); Key Largo, under bark, 29-VIII-1961, H.V. Weems, Jr. (1, FSCA); Key Largo, 15-V-

1977, 27-V-1983, R. Turnbow, 12-V-1975, E. Giesbert (9, FSCA; 3, RHTC); upper Key Largo, at light, 9-VI-1995, M.C. Thomas (4, FSCA); upper Key Largo, 3-VI-1993, R. Morris and

Bratain, 7-VI-1998, R. Morris, 7-VI-1976, R. Turnbow (3, FSCA); upper Matecumbe Key, 1 mile southwest of Islamorada, blacklight trap, 27-IV-1975, J.B. Heppner (1, FSCA); Orange

County: Rk Spr Rn St Res, Spine/Oak Scrub, Malaise Trap, 14-VI-1995, S.M. Fullerton (1,

UCFC); University of Central Florida, Orlando, LLP-Sand Pine, Turkey Oak, Malaise Trap, 11-

VI-1997, S.M. Fullerton (1, UCFC); Orlando, alpha-beta pinene baited Lindgren funnel trap, 27-

IV-2007, 4-VI-2007, B. Saunders (7, CAPS); Orlando, ethanol-baited Lindgren funnel trap, 30-

III-2007, 27-IV-2007, 6-VI-2007, 18-IX-2006, 11-X-2006, B. Saunders, VI-2007, J.M.

Leavengood, Jr. (11, CAPS; 4, JMLC); Orlando, McKay Tract Apartment Complex, edge red maple hydric hammock, at UV light, 2-III-2000, T. Smith (1, FSCA); Pinellas County: Saint

Petersburg, ethanol-baited Lindgren funnel trap, IV-2007, VI-2007, J.M. Leavengood, Jr. (5,

JMLC); Santa Rosa County: 1.7 miles north of Munson, 20-IV-1991, R. Turnbow (1, RHTC); 4 miles north of Munson, at mercury vapor and black light, 7-VI-1986, R. Turnbow (1, RHTC);

Sarasota County: Englewood, residential/urban disturbed yard, blacklight trap, 26-III-1997, K.

Maharay and S.M. Fullerton (2, UCFC); Seminole County: Oviedo, rural yard, ultraviolet light trap, 27-IV-1994, S.M. Fullerton (2, UCFC); Sumter County: I-75 and CR 470, 18-IV-1992, R.

Turnbow (1, FSCA); Taylor County: 13 miles north of Steinhatchee, on CR 361, 13-V-1995,

M.C. Thomas (1, FSCA); Volusia County: Daytona, IV-1952, W. Rosenberg (2, FSCA); VIII-

1956, H.A. Denmark (1, FSCA); Wakulla County: 6 miles east of Newport, 4-VI-1988, R.

Turnbow (1, RHTC); Walton County: Highway 285 at Shoal River, 5-V-1985, R. Turnbow (1,

RHTC); Washington County: Falling Waters State Park, 29-VI-1970, G.H. Nelson (1, FSCA).

Selected references: Fabricius, 1798: 177; Say, 1823: 188; Dejean, 1837: 128; Klug, 1842: 358; Chevrolat, 1843: 34; Spinola, 1844: 80, 82, pl. 48, fig. 4; Melsheimer, 1846: 307; LeConte, 1849: 33, 1859a: 90; Chevrolat, 1874: 325; Horn, 1875: 149; Drury, 1879: 171; Gorham, 1883: 185, 1886: 345; Hopkins, 1893a: 186; Horn, 1894: 331; Hamilton, 1895: 335; Wickham, 1895: 252; Lohde, 1900: 109; Schenkling, 1903a: 110, 1906: 317; Wolcott, 1909: 101, 1910b: 395, 1910c: 861; Chapin, 1920: 52; Wickham and Wolcott, 1912: 67; Bøving and Champlain, 1920: 618, 639; Knull, 1930: 83; Bøving and Craighead, 1931b: 276; Knull, 1932: 43; Wolcott, 1943: 137; Beal and Massey, 1945: 122; Hoffmann, 1942: 7; Wolcott, 1947: 84; Barr, 1950a: 512; Corporaal, 1950: 268; Craighead, 1950: 203; Knull, 1951: 304, pl. 10, fig. 40; Vaurie, 1952a: 4; Papp, 1960: 85; Ekis, 1975: 63; Barr, 1976b: 7; Barr, 1975: 15; Gosling, 1980: 74; Peck and Thomas, 1998: 86; Opitz, 2002: figs. 85, 104, 105.

Genus Pelonium Spinola, 1844

= Corinthiscus Fairmaire and Germain, 1861

Pelonium is represented by four widely distributed species in the United States (Opitz,

2002: 279). This genus is distinguished by its emarginated eyes, 11-segmented antennae and angulate sides of the pronotum which is constricted basally. Most obvious in the Florida fauna is the lichen-mimicking color patterns.

Selected references: Spinola, 1844: 347; Fairmaire and Germain, 1861: 4; Wolcott, 1910b: 383, 1947: 86; Opitz, 2002: 279.

Pelonium leucophaeum (Klug), 1842 (Figs. 3-11, 3-12)

= Pelonium vestutum Spinola, 1844 = Priocera albomaculata Zeigler, 1845

Pelonium leucophaeum is distinguished by its lichen-mimicking color pattern and pronotum with irregularly elevated regions. The elytra are irregularly punctured. The body is clothed with semierect pale hairs. Adults are 6 to 11 mm long.

Pelonium granulosum Wolcott is strikingly similar. However, P. granulosum has

(sometimes subtly) less elevated regions on the pronotum and has not been recorded from

Florida.

Biology: Specimens have been reared from small trees and branches of cypress and

juniper infested with immature Cerambycidae (Coleoptera) (Bøving and Champlain, 1920: 638;

Opitz, 2002: 279). It has been collected on poison ivy (Wolcott, 1910b: 385).

Specimens have been collected in Florida from February through September. They have

been collected in Malaise traps, Steiner traps, Mediterranean fruit fly traps, suction traps, at

ultraviolet lights, and in Lindgren funnel traps baited with ethanol and alpha-beta pinene

(separately). Adults have been reared from Taxodium distichum (Linnaeus) (Taxodiaceae) and

have been collected on Citrus aurantium (Linnaeus) (Rutaceae), and Sambucus canadensis

Linnaeus (Caprifoliaceae).

Distribution: Alabama, California, Florida, District of Columbia, Illinois, Kansas,

Louisiana, Michigan, Missouri, New Jersey, New York, Ohio, Pennsylvania, South Carolina,

Texas and Mexico (Wolcott, 1909: 100, 1910b: 385, 1947: 86; Knull, 1951: 308; Papp, 1960: 86;

Gosling, 1980: 74; Barr, 1975: 17). Wickham and Wolcott (1912: 66) recorded it from

Allegheny County, Pennsylvania, which is within 50 miles of Maryland and West Virginia.

In Florida it has been recorded from Alachua, Bay, Brevard, Broward, Duval, Flagler,

Hernando, Highlands, Indian River, Lake, Liberty, Manatee, Marion, Miami-Dade, Orange,

Pinellas, Polk, Putnam, Saint John, Santa Rosa, Sarasota, and Wakulla Counties. Wolcott (1910:

385) recorded it from Enterprise, Volusia County, Florida.

Specimens examined (93): FLORIDA: Alachua County: Gainesville, 29-V-1964, R.E.

White, 23-VI-1977, R.A. Belmont, 30-II-1984, M.C. Thomas, 1-VII-1956, T. Denmark (4,

FSCA); SE Gainesville, Kincaid Rd, old field edge, xeric oak habitat, 6 meter Malaise trap, 24-

IV to 1-V-1999, B.D. Sutton (1, FSCA); 18-VII-1958, T. Denmark (1, FSCA); Bay County: ethanol-baited Lindgren funnel trap, 13-VI-2007, L. Smith (1, CAPS); Brevard County:

Titusville, SR 405, Enchanted Forest Sanctuary, White Trail, xeric oak hammock, Malaise Trap,

21-II-7-III-2001, P. Russell, Z. Prusak and S.M. Fullerton (1, UCFC); Broward County:

Hollywood, on Citrus paradisi, 4-V-1994, B. Coy (1, FSCA); Duval County: Jacksonville,

ethanol-baited Lindgren funnel trap, 7-VI-2007, A. Silagyi and A. Johnson, 18-V-2007, A.

Johnson (3, CAPS); Flagler/Saint John's County: Marineland, in Steiner trap, 28-V-1962, G.W.

Desin (1, FSCA); Hernando County: Withlacoochee State Forest, ex. Taxodium distichum (L.), emerged 26-IV-1991, R. Turnbow (10, RHTC); Withlacoochee State Forest, Goat Road, 28-IV-

1990, R. Morris (1, FSCA); Withlacoochee State Forest, Richloam Unit, 27-IV-1991, R.

Turnbow (1, RHTC); Highlands County: Archbold Biological Research Station, at ultraviolet light, 16-IV-1978, 26-IV-1976, L.L. Lampert, Jr. (2, FSCA); Indian River County: Fellsmere, on

Sambucus canadensis, 18-IV-1983, E.W. Campbell (1, FSCA); SR 512, 0.5 miles west of I-95, suction trap, 19-VI-1975, (1, FSCA); Vero Beach, 25-V-1964, 8-VI-1964, W.L. Bidlingmayer

(3, FSCA); Lake County: Clermont, in Steiner trap, 21-VI-1966, C.L. Felshaw (1, FSCA);

Mascotte, in medfly trap, VII-1960, C.L. Felshaw (1, FSCA); Liberty County: Apalachicola

National Forest, ethanol-baited Lindgren funnel trap, V-2007, J.M. Leavengood, Jr. (1, JMLC);

Torreya State Park, 9-VII-1988, R. Turnbow (1, RHTC); Manatee County: Palma Sola, in

Steiner trap, 27-V-1963, D.C. Chancey (1, FSCA); Marion County: McIntosh, ethanol-baited

Lindgren funnel trap, IV-2007, J.M. Leavengood, Jr. (3, JMLC); Miami-Dade County: Ojus, in

Steiner trap, 29-IV-1971, D.R. Dehaven (1, FSCA); 20-IV-1949, (1, FSCA); Orange County:

Orlando, alpha-beta pinene-baited Lindgren funnel trap, 4-VI-2007, B. Saunders (1, CAPS);

Orlando, ethanol-baited Lindgren funnel trap, 27-IV-2007, 6-VI-2007, B. Saunders, VI-2007,

J.M. Leavengood, Jr. (4, CAPS; 5, JMLC); Orlando, 20-V-1964, J.R. Woodley (1, FSCA); Rk

Spr Rn St Res, S Pine/Oak Scrub, Malaise Trap, 27-IV-1995, S.M. Fullerton (1, UCFC);

University of Central Florida, MacKay Tract, Orlando, Sawgrass Marsh, red maple, Malaise

Trap, 24-V-1999, P. Russell and S.M. Fullerton (1, UCFC); University of Central Florida,

Orlando, Cypress Dome, Malaise Trap, 20-V-1993, S.M. Fullerton (1, UCFC); University of

Central Florida, Orlando, Cypress Forest, Malaise Trap, 24-V-1999, 8-VI-1999, 2-VI-1999, 30-

VIII-1999, P. Russell and S.M. Fullerton (5, UCFC); University of Central Florida, Orlando,

LLP-Sand Pine, Turkey Oak, Malaise Trap, 2-VII-1997, 21-VIII-1993, S.M. Fullerton (2,

UCFC); University of Central Florida, Orlando, Longleaf Pine-Saw Palmetto, Malaise Trap, 11-

VI-1997, S.M. Fullerton (1, UCFC); University of Central Florida, Orlando, Maidencane Marsh,

Malaise Trap, 2-VI-1999, 8-VI-1999, 17-V-1999, 24-V-1999, 19-IV-1999, 26-IV-1999, 11-V-

1999, 5-IV-1999, 14-VI-1999, P. Russell and S.M. Fullerton (14, UCFC); University of Central

Florida, Orlando, Pond Pine Community, Dahoon Holly, Malaise Trap, 2-VI-1999, 8-VI-1999,

17-V-1999, 2-VIII-1999, P. Russell and S.M. Fullerton (4, UCFC); Pinellas County: Fort Desoto

State Park, 3-VI-1998, R. Morris (1, FSCA); Saint Petersburg, ethanol-baited Lindgren funnel trap, VI-2007, J.M. Leavengood, Jr. (1, JMLC); Polk County: Frostproof, in Steiner trap, 25-IV-

1961, R.E. Vild (1, FSCA); Putnam County: Lake Elizabeth, ex. Taxodium sp., emerged 10-V-

2000, R. Turnbow (1, RHTC); Santa Rosa County: 11.7 miles east of Gulf Breeze, 18-VI-1988,

R. Turnbow (1, RHTC); Gulf Breeze, ethanol-baited Lindgren funnel trap, 4-V-2007, 24-V-

2007, 14-IX-2006, C. Street (3, CAPS; 1, JMLC); Sarasota County: Sarasota, on Citrus aurantium, 3-VI-1987, K. Jenkins (1, FSCA); Wakulla County: 6 miles east of Newport, 4-VI-

1988, R. Turnbow (2, RHTC); [county unknown]: Parish, 29-VII-1964, D.C. Chancey (1,

FSCA);

Selected references: Klug, 1842; 366; Spinola, 1844: 360, pl. 35, fig. 4; Zeigler, 1845: 268; Haldeman, 1847: 126; LeConte, 1849: 32; Melsheimer, 1853: 84; LeConte, 1865: 98; Chevrolat, 1876: 7; Henshaw, 1885: 82; Hamilton, 1895: 335; Horn, 1895: 228; Lohde, 1900: 103; Schenkling, 1903a: 107, 1903b: 16; Schaeffer, 1904: 220; Schenkling, 1906:

316, 1907: 299; Wolcott, 1909: 100, 1910b: 385, 1910c: 860; Wickham and Wolcott, 1912: 66; Bøving and Champlain, 1920: 638; Wolcott, 1947: 86; Knull, 1951: 307, pl. 11, fig. 44; Papp, 1960: 86; Ekis, 1975: 57; Barr, 1975: 17; Gosling, 1980: 74; Opitz, 2002: 279, figs. 76, 78, 84, 92, 115.

Figure 3-1. Chariessa floridana

Figure 3-2. Distribution map of Chariessa floridana

Figure 3-3. Chariessa pilosa

Figure 3-4. Distribution map of Chariessa pilosa

Figure 3-5. Cregya oculata

Figure 3-6. Distribution map of Cregya oculata

Figure 3-7. Cregya quadrinotata

Figure 3-8. Distribution map of Cregya quadrinotata

Figure 3-9. Neorthopleura thoracica

Figure 3-10. Distribution map of Neorthopleura thoracica

Figure 3-11. Pelonium leucophaeum

Figure 3-12. Distribution map of Pelonium leucophaeum

CHAPTER 4 EPIPHLOEINAE

Key to the Florida Genera and Species of Epiphloeinae

1. Antennal club shorter than funicle; pronotum entirely brown; elytra patterned ...... Madoniella dislocatus (Fig. 4-1)

1’. Antennal club longer than funicle; pronotum orange, with a longitudinal black vitta; elytra entirely black ...... Pyticeroides laticornis (Fig. 4-3)

Genus Madoniella Pic, 1935

This genus is represented by three species across the eastern, central and southern United

States, and Texas to California (Barr 1962: 126). Other members of the genus occur in the West

Indies and South America. Madoniella is currently being revised by Weston Opitz, and includes numerous new species (personal communication). Opitz (1997: 62) redescribed the genus.

They are small and elongate, with anteriorly emarginate eyes, ten-segmented antennae with a three-segmented club which is shorter than the funicle, a subquadrate pronotum, and slender tarsi with strongly dentate tarsal claws. The antennal club of Pyticeroides is longer than the funicle, distinguishing it from Madoniella.

Bøving and Champlain (1920: 609) described the larvae and compared them to some related taxa.

Selected references: Spinola, 1844: 1; LeConte, 1849: 30; Lacordaire, 1857: 466; Desmarest, 1860: 264; Gemminger and Harold, 1869: 1747; Gorham, 1877: 246; LeConte and Horn, 1883: 219; Kuwert, 1893: 492; Wickham, 1895: 248; Lohde, 1900: 87; Schenkling, 1903a: 86, 1910: 113; Wolcott, 1910c: 858; Bøving and Champlain, 1920: 609; Gahan, 1910: 59, 73; Bradley, 1930: 107; Bøving and Craighead, 1931a: 57; Pic, 1935: 100; Wolcott, 1944a: 124, 1947: 83; Fleutiaux, 1947: 169; Winkler, 1961: 59; Barr, 1962: 126; Dillon and Dillon, 1972: 279; Opitz, 1997: 62, figs. 109-120, 2002: 278.

Madoniella dislocatus (Say), 1825 (Figs. 4-1, 4-2)

= Enoplium distrophum Klug, 1842 = Phyllobaenus transversalis Spinola, 1844

Madoniella dislocatus has yellow to light brown mouth parts, antennae and legs. The pronotum is entirely brown and the elytra are patterned with dark brown and yellow. The head,

pronotum and elytra are densely, coarsely punctate. The posterior elytral margins are serrulate.

Adults are 3.5 to 6.0 mm long. The generic characters and color patterns distinguish Madoniella

dislocatus from Pyticeroides laticornis.

Bøving and Champlain (1920: 610) described the larvae and compared them to some related taxa.

Biology: Madoniella dislocatus is commonly found in the burrows of many wood-boring insects. Their most common prey are bark beetles associated with oaks, juniper, cedar, hickory, pine, hackberry, black spruce, sumac, larch, butternut, redbud and other hardwoods (Opitz, 2002:

278). Adults are diurnal and can be found on dead branches of trees, especially on oak between

May and August. They have been recorded from oak, red oak, hickory, Rhus sp.

(Anacardiaceae), redbud, hackberry, apple, and black spruce infested with Elaphidion villosum

(Fabricius) (Coleoptera: Cerambycidae), Bitoma carinata (LeConte) (Colydiidae), Agrilus lecontei Saunders (Buprestidae), Chramesus hicoriae LeConte, Micrasis sp., Scolytus rugulosus

(Muller), Scolytus muticus Say, Phloeotribus dentifrons Blackman, Polygraphus rufipennis

(Kirby) and Pityophthorus consimilis LeConte (Coleoptera: Curculionidae: Scolytinae) (Knull,

1951: 302; Gosling, 1980: 73; Mawdsley, 1999: 44).

Adults have been collected in Florida from March through May. Specimens have been collected in ethanol-baited Lindgren funnel traps and sticky traps.

Distribution: Colorado, District of Columbia, Florida, Georgia, Indiana, Illinois, Iowa,

Kansas, Maine, Massachusetts, Michigan, New Hampshire, New Jersey, New York, Ohio,

Pennsylvania, South Carolina, Texas, West Virginia, Wisconsin and Ontario (Wickham and

Wolcott, 1912: 65; Wolcott, 1947: 84; Barr, 1975: 14). In Florida it has been recorded from

Alachua, Jackson, Leon, Liberty, and Union Counties.

Specimen examined (19): FLORIDA: Alachua County: Archer, sticky wire trap, III-

1984, M.J. Plagens (1, FSCA); Jackson County: Florida Caverns State Park, 3-5-V-2008, M.C. and S. Thomas (13, FSCA); Leon County: Tallahassee, ethanol-baited Lindgren funnel trap, 4-V-

2007, M. Bentley (1, CAPS); Liberty County: Torreya State Park, 27-IV-1991, M.C. Thomas,

16-V-1964, R.E. White (2, FSCA); Torreya State Park, ethanol-baited Lindgren funnel trap, V-

2007, J.M. Leavengood, Jr. (1, JMLC); Union County: Highway 238-A, 4-IV-1989, C.W. Mills,

III (1, FSCA).

Selected references: Say, 1825: 176; Klug, 1842: 374; Spinola, 1844: 4, pl. 40, fig. 6; LeConte, 1849: 30, 1859a: 284; Drury, 1879: 171; Schwarz, 1890: 167; Hopkins, 1893a: 156; Hamilton, 1895: 335; Lohde, 1900: 87; Felt, 1906: 503; Wolcott, 1909: 99, 1910c: 858; Wickham and Wolcott, 1912: 65; Bøving and Champlain, 1920: 610, 636; Blackman and Stage, 1924: 45; Beal and Massey, 1945: 69, 71, 78, 79; Wolcott, 1947: 84; Knull, 1951: 302, pl. 10, fig. 39; Dillon and Dillon, 1972: 280, pl. 29, fig. 1; Ekis, 1975: 46; Barr, 1975: 14; Gosling, 1980: 73; Opitz, 1997: figs. 109-120; Peck and Thomas, 1998: 86; Mawdsley, 1999: 44; Opitz, 2002: 278, figs. 75, 95.

Genus Pyticeroides Kuwert, 1894

Pyticeroides is represented in North American by a single species, Pyticeroides laticornis

(Say), which is distributed in the eastern and central United States (Barr 1962: 126). Other members of the genus are distributed across Central and South America.

Pyticeroides is characterized by having nine antennomeres. Other epiphloeines have 8, 10 or 11 antennomeres. The three-segmented antennal club is longer than the funicle. In

Madoniella the antennal club is shorter than the funicle.

Opitz (1997: 59, 2007: 89) redescribed Pyticeroides, which was previously known to have only five species and all but two were of different genera (Ellipotoma, Apolopha, Enoplium and

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Pyticeroides). His revision added 27 new species, making the genus comprehensively defined

and understood for the first time in history.

Opitz (2007: figs. 123-127) illustrated several larval structures of the genus and a habitus

illustration of a P. laticornis larva. These illustrations were not accompanied by a diagnosis or

discussion. Bøving and Champlain (1920: 610) described the larvae and compared them to some

related taxa.

Selected references: Castelnau, 1836: 55; LeConte, 1849: 31; LeConte and Horn, 1883: 219; Kuwert, 1894: 7, 9; Wickham, 1895: 248; Lohde, 1900: 103; Schenkling, 1903a: 112; Gahan, 1910: 73; Wolcott, 1910c: 858; Wolcott and Chapin, 1918: 108; Bøving and Champlain, 1920: 610; Chapin, 1927: 8; Bradley, 1930: 107; Bøving and Craighead, 1931a: 57; Corporaal, 1942: 141; Blackwelder, 1945: 388; Wolcott, 1947: 84; Corporaal, 1950: 253; Winkler, 1961: 59; Barr, 1962: 126; Opitz, 1997: 59, figs. 78, 79, 80, 2002: 278, 2007: 87.

Pyticeroides laticornis (Say), 1835 (Figs. 4-3, 4-4)

This elongate species has black, patternless elytra that are over six times longer than either the pronotal length or elytral width. The head and pronotum are orange to yellow, with black vittae extending from the posterior margins of the eyes to the pronotum, where they fuse into a single broad vitta spanning the length of the pronotum. The head and pronotum are densely, finely punctate. Adults are about 6.0 mm long.

The most notable color variation is the pale recumbent setae on the head and lateral pronotum. In some specimens these setae are gold to yellow, in others they are more cream colored (Opitz, 2007: 131). The generic characters and color pattern distinguish it from

Madoniella dislocatus.

Biology: This diurnal species has been recorded from elm, red cedar, oak, hickory, eastern redbud, mulberry, juniper, hackberry, black cherry, and cypress infested with Chramesus hicoriae LeConte, Scolytus quadrispinosus Say, Scolytus multistriatus (Marsham), Phloeotribus liminaris (Harris), Phloeophthorus sp. and Phloeosinus taxodii Blackman (Coleoptera:

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Curculionidae: Scolytinae) (Bøving and Champlain, 1920: 637; Hoffman, 1940: 56; Knull, 1951:

303; Gosling, 1980: 73; Opitz, 2002: 278). The adults are diurnal and have been observed feeding on scolytid eggs and larvae in galleries (Bøving and Champlain, 1920: 637). At

infestation sites, the adult female lays a single egg at the gallery entrance of a bark beetle. The

larva preys upon scolytid adults, larvae and eggs. It has been collected from February to October

by beating, trapping or at ultraviolet light (Opitz, 2007: 132).

Mawdsley (1994: 116) considers P. laticornis to be a part of a tremendous coleopteran

mimicry ring with Lampyridae, Cantheridae, Lycidae, Buprestidae, Meloidae, Ptilodactylidae,

Elateridae, and the Cleridae Placopterus thoracicus, Monophylla terminata and Chariessa

pilosa.

Adults have been collected in Florida in January, February, April and May.

Distribution: Alabama, Arkansas, Colorado, Connecticut, District of Columbia, Florida,

Georgia, Illinois, Indiana, Kansas, Kentucky, Maine, Maryland, Massachusetts, Michigan,

Missouri, Mississippi, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania,

South Carolina, Tennessee, Texas, Virginia, West Virginia and Ontario (Wickham and Wolcott,

1912: 66; Wolcott, 1947: 84; Knull, 1951: 304; Papp, 1960: 85; Barr, 1975: 15; Gosling, 1980:

73). In his revision of the genus Pyticeroides, Opitz (2007: 131) suggested Mexico as part of

this distribution because some Mexican specimens examined “do not differ significantly from P.

laticornis”.

In Florida P. laticornis has been recorded from Alachua, Jackson and Orange Counties.

Opitz (2007: 131) recorded specimens from Orlando, Orange County, 31-I-1997, ex. Scolytid

infested Leyland Cypress, and Archer, Alachua County, “Guin” SV, W. S. B. Mariana, W. F.

Fiske, reared 14-IV-1905.

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Specimens examined (2): FLORIDA: Alachua County: Archer, sticky wire trap, II-1984,

M.J. Plagens (1, FSCA); Jackson County: Florida Caverns State Park, 3-5-V-2008, M.C. and S.

Thomas (1, FSCA).

Selected references: Say, 1835: 164; LeConte, 1849: 31, 1859b: 641; Hamilton, 1895: 335; Lohde, 1900: 101; Drury, 1902: 149; Wolcott, 1910c: 859; Wickham and Wolcott, 1912: 66; Bøving and Champlain, 1920: 611, 637; Blackman and Stage, 1924: 45; Bøving and Craighead, 1931b: 276; Hoffmann, 1940: 56, 1942: 7; Beal and Massey, 1945: 76, 79, 87; Wolcott, 1947: 84; Craighead, 1950: 203; Knull, 1951: 303, pl. 10, fig. 41; Papp, 1960: 85; Barr, 1975: 15; Gosling, 1980: 73; Mawdsley, 1994: 116; Opitz, 1997: figs. 78-90, 2002: 278, figs. 1, 11, 79, 2007: 130, figs. 27-29, 39-46, 49-51, 56, 66, 95, 116, 123-127.

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Figure 4-1. Madoniella dislocatus

Figure 4-2. Distribution map from Madoniella dislocatus

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Figure 4-3. Pyticeroides laticornis

Figure 4-4. Distribution map of Pyticeroides laticornis

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CHAPTER 5 HYDNOCERINAE

Key to the Florida Genera of Hydnocerinae

1. Elytra either entirely blue to green or patterned, never entirely pale testaceous .... Phyllobaenus

1’. Body entirely pale testaceous, occasionally with a black macula on the head ...... Isohydnocera

Genus Isohydnocera Chapin, 1917

This genus is represented by 14 species throughout the eastern, central and southern United

States to Arizona (Barr, 1962: 125). Other members of the genus occur throughout the Western

Hemisphere.

According to current literature Isohydnocera is distinguished by its simple tarsal claws.

The tarsal claws of Phyllobaenus are bifid. Previous literature has indicated that the third antennomere in Isohydnocera is twice as long as it is broad, and the elongate bodies and spindly, delicate legs help separate them from Phyllobaenus. There is much overlap in these characters which previous authors considered distinctive. Additionally, recent authors diagnosed

Isohydnocera citing characters that disagree with the original descriptions of the species discussed within their publication (Knull, 1951). Because specimens identified as I. aegra do not match the published description, I. aegra is considered a synonym in this work. It is considered that I. aegra does not exist (Barr via Thomas, personal communication). Much work is needed to confidently distinguish Isohydnocera from Phyllobaenus.

Mawdsley (1994: 117) considers the members of the genus to be generalized ant mimics.

Selected references: Chapin, 1917: 83; Bradley, 1930: 105; Wolcott, 1944a: 127, 1947: 74; Barr, 1962: 125; Mawdsley, 1994: 117; Opitz, 2002: 276.

Isohydnocera curtipennis (Newman), 1840 (Figs. 5-1, 5-2)

= Hydnocera longicollis Zeigler, 1844 = Isohydnocera aegra (Newman), 1840

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Isohydnocera curtipennis is entirely pale testaceous. Previous literature suggested that the

elytra and or head and prothorax were patterned (Knull, 1951: 283). The body is elongate and

narrow with delicate, spindly legs. It resembles an elongated, patternless version of

Phyllobaenus. Adults are about 5.0 mm long. Sabrosky (1934: 66) described the larva,

discussed its similarity to other clerid taxa, and explained how to use the work of Bøving and

Champlain (1920) regarding this species.

There is some confusion as to whether past literature referred to I. curtipennis or a member of Phyllobaenus (see the discussion under Isohydnocera).

Biology: Isohydnocera curtipennis has been considered to breed in hickory and witch hazel as a borer or predator of wood-boring insect larvae. It has been recorded from galls of

Euura salicus-nodus Walsh (Hymenoptera: Tenthredinidae) on willow and Gnorimoschema

gallaesolidaginis Riley (Lepidoptera: Gelechiidae) on golden rod (Sabrosky, 1934: 65; Knull,

1951: 284). Sabrosky (1934: 65), after finding the exuvium of the beetle inside of the pupal cell,

suggested the immatures consume the contents of solidago gallmoth pupae while occupying the

pupal cell as an endoparasite. Isohydnocera curtipennis has been captured in a habitat rich with

torpedograss on Lake Okeechobee (Cuda, 2007: 243).

In Florida adults have been collected March through May. One specimen was collected in

a Malaise trap.

Distribution: Connecticut, District of Columbia, Florida, Georgia, Illinois, Indiana,

Kansas, Maryland, Michigan, Minnesota, Missouri, New York, North Carolina, Ohio,

Pennsylvania, South Carolina, Wisconsin, and Ontario (Newman, 1840: 364; Zeigler, 1844: 44;

Sabrosky, 1934: 65; Wolcott, 1947: 74; Papp, 1960: 80; Barr, 1975: 14; Gosling, 1980: 72).

Knull (1951: 284) recorded it from Ohio, stating that adults occur from May through August.

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In Florida it has been recorded from Alachua, Collier, Dixie, Franklin, Gulf, Indian River,

Liberty, Palm Beach, Sarasota, Saint Lucie, and Santa Rosa Counties. Cuda (2007: 243) recorded it from Glades County. Wickham and Wolcott (1912: 65) recorded it from Crescent

City, Putnam County.

Specimens examined (2): Florida: Liberty County: Torreya State Park, 4-V-2007, J.M.

Leavengood, Jr. (1, JMLC); St. Lucie County: Fort Pierce, UF/IRREC, Malaise trap, 12-III-

2007, C.L. Branch (1, JMLC).

Selected references: Newman, 1840: 364; Zeigler, 1844: 44; LeConte, 1849: 29; Drury, 1879: 171, Chittenden, 1890: 155; Baker, 1895: 28; Hamilton, 1895: 335; Lohde, 1900: 92; Wolcott, 1909: 100, 1910c: 856; Wickham and Wolcott, 1912: 65; Sabrosky, 1934: 65; Wolcott, 1947: 74; Knull, 1951: 283, pl. 4, fig. 13; Papp, 1960: 80; Barr, 1975: 14; Gosling, 1980: 72; Peck and Thomas, 1998: 86; Mawdsley, 1999: 44; Opitz, 2002: 276; Mawdsley, 2002b: 20; Cuda, 2007: 243.

Genus Phyllobaenus Dejean, 1833

Phyllobaenus is represented by 58 well-distributed species, seven of which occur in

Florida (Barr, 1962: 125). Members of this genus are often about 4.0 mm long, with soft elytra which often do not cover the entire abdomen, entire, finely granulate eyes, and a short hind body.

The posterior elytral margins are minutely serrulate and the anteapical region of each elytron is tumid in some species. The tarsal claws are broadly dentate, the maxillary palpi are cylindrical with the apical palpomeres weakly tapering, and the labial palpi are elongate with the apical palpomere strongly dilated. The antennae are nearly filiform with an abruptly formed, compact club of two subglobose segments.

Phyllobaenus, Isohydnocera and Wolcottia are the only genera of the subfamily in the

United States. All genera in the subfamily share varying degrees of the posterior elytral serrulation and body form. Separating members of Phyllobaenus from Isohydnocera requires revisionary work (see the discussion of Isohydnocera). Most members of Phyllobaenus are

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smaller, have more robust bodies, and basally toothed tarsal claws which separate them from

Isohydnocera. Wolcottia is similar in body form to Phyllobaenus and has a tooth-like notch on its tarsi, where Phyllobaenus has much more distinct tarsal denticles.

Wolcott (1944a: 123) offered an extensive discussion of the nomenclatural changes throughout the literary history of the genus with an updated genus diagnosis. Bøving and

Champlain (1920: 622) described the larvae of “Hydnocera” and compared them to some related taxa. At the time of publication, Hydnocera included Phyllobaenus and Isohydnocera.

However, because the work only addressed species now known as Phyllobaenus, it should only be considered a genus treatment of Phyllobaenus.

Members of Phyllobaenus are often collected in sweep samples of grassy and short foliage habitats. They have been reared from cotton bolls and insect galls. Other plant associations include wild grape, river birch, black and white oak, hickory, sumac, pine, blackberry, cotton, elm, and desert shrubs. These beetles are predacious on small wood-boring insects, wasp larvae and immature weevils (Opitz, 2002: 276).

Mawdsley (1994: 117) considers all members of Phyllobaenus to be generalized ant mimics. Some species are also chemical mimics of Pheidole ants (Hymenoptera: Formicidae), producing chemicals which induce plant tissue growth in stems.

My observations suggest a need for revisionary work to accurately distinguish many species of Phyllobaenus. The problem lies within a group of species here named the pallipennis- verticalis complex, including the known species P. pallipennis, P. suturalis, and P. verticalis. It is uncertain if P. suturalis is a part of this complex or if it even occurs in Florida. This is due to extensive character variation seen in members of these ‘species’ and the all too brief, old (1825-

1842) original species description, to which recent literature has added little usable data. As

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such, P. suturalis has been excluded from the diagnostic key (below) but included as a species

treatment. I also consider that a species, or variation of a described species, of Phyllobaenus is

commonly mistakenly considered to be a member of Isohydnocera (see the discussion of

Isohydnocera). The key below is written according to the most current literature and should

work for most specimens encountered.

Selected references: Dejean, 1833: 113; Newman, 1838: 379; Chevrolat, 1843: 33; LeConte, 1849: 26; LeConte and Horn, 1883: 218; Wolcott, 1910c: 855; Chapin, 1917: 83; Bøving and Champlain, 1920: 622, 635; Bradley, 1930: 105; Wolcott, 1944a: 123, 1947: 65, 71; Barr, 1962: 125; Dillon and Dillon, 1972: 274; Mawdsley, 1994: 117; Opitz, 2002: 276.

Key to the Florida Species of Phyllobaenus

1. Legs entirely pale yellow to orange ...... 2

1’. Legs bicolored or entirely bluish-black to black ...... 4

2 (1). Elytra somewhat metallic bluish to green, but unicolorous; pronotum matching elytral color, lacking any maculations; head matching elytral color ...... P. rufipes (Fig. 5-9)

2’. Elytra black, with yellowish maculations of variable form; pronotum black, with or without yellowish maculations; head yellowish or black ...... 3

3 (2). Pronotum lacking any yellowish maculations; head black, with yellow integument only at mouthparts and antennae; elytra with four yellowish maculae of variable size, sometimes covering most of the elytral surface ...... P. pallipennis (Fig. 5-5)

3’. Pronotum variable, with or without yellowish maculations; head yellow, with or without a black macula located basally between the eyes; elytra with two humeral maculae which may extend apically to the full length of the elytra covering most of the surface, if so extended, some evidence of black should remain at the elytral apices and/or elytral margins ...... P. verticalis (Fig. 5-15)

4 (1). Head, pronotum and elytra unicolorous, lacking any maculations; body covered with dense pubescence; legs orangish red ...... P. pubescens (Fig. 5-7)

4’. Head, pronotum and elytra bluish black to black, but with two elytral maculae; body lacking dense pubescence; legs black and unicolorous or bluish black with orangish red at the base of the femora and first tarsomeres ...... 5

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5 (4). Elytra with orangish humeral maculae; legs bluish black with orangish red at the base of the femora and first tarsomeres ...... P. humeralis (Fig. 5-3)

5’. Elytra with pale median transverse fascia; legs black and unicolorous ...... P. unifasciatus (Fig. 5-13)

Phyllobaenus humeralis (Say), 1823 (Figs. 5-3, 5-4)

= Phyllobaenus axillaris Dejean, 1833

Phyllobaenus humeralis is a somewhat robust, dark bluish-black hydnocerine best recognized by the orange spots on its elytral humeri. Other than the humeral spots, its body, legs, pronotum and head are bluish-black. The mouthparts, antennae and parts of the legs are yellowish to brown. The yellowish on the legs may be found often on the apical profemora and protibiae and sometimes on the mesofemora. There is much variation and some specimens have legs that are nearly entirely bluish black. The margins of the apical third of the elytra are weakly and rather bluntly serrulate. If the serrulations appear pointed, then the length of the serrulation should not exceed the width of nearby elytral punctures. The apical third of the elytra is convex.

Adults range from 3.5 to 5.5 mm in length.

Phyllobaenus humeralis resembles P. pubescens and P. rufipes, both of which lack

humeral spots and have uniformly reddish-orange legs. While Phyllobaenus humeralis is similar

in pattern, Necrobia ruficollis is a metallic greenish-blue clerid with orangish humeri, pronotum,

and legs. But, N. ruficollis differs significantly in body form and generic characters.

Biology: Mawdsley (2002b: 21) noted that P. humeralis breeds in the trunks and limbs of

oaks and hickory, and that it may also occur on flowers and foliage of herbaceous plants.

A single specimen collected in Florida in April.

Distribution: Arizona, California, Colorado, District of Columbia, Florida, Georgia,

Idaho, Illinois, Indiana, Kansas, Kentucky, Maine, Maryland, Massachusetts, Michigan,

Minnesota, Missouri, Montana, Nebraska, New Hampshire, New Jersey, New Mexico, New

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York, North Carolina, North Dakota, Ohio, Oregon, Pennsylvania, South Carolina, South

Dakota, Texas, Utah, Virginia, Washington, Wisconsin, Mexico, British Columbia, Manitoba,

Ottawa and Ontario (Wolcott, 1909: 99; Wickham and Wolcott, 1912: 61; Wolcott, 1947: 72;

Knull, 1951: 281; Papp, 1960:79; Barr, 1975: 12). This distribution includes the subspecies P. h. cyaneus and P. h. difficilis, however only P. h. humeralis is known from Florida.

In Florida only one specimen from Wakulla County has been collected. Wickham and

Wolcott (1912: 61) and Papp (1960: 79) recorded this species from Florida, although neither reference referred directly to a specimen.

Specimens examined (1): FLORIDA: Wakulla County: Ochlockonee State Park, 4-IV-

1984, N.M. Downie (1, FSCA).

Selected references: Say, 1823: 192; Dejean, 1833: 133; Newman, 1840: 362; Spinola, 1844: 44; LeConte, 1849: 27, 1859b: 122, 640; Hamilton, 1895: 335; Lohde, 1900: 93; Drury, 1902: 149; Wolcott, 1909: 99, 1910c: 856; Wickham and Wolcott, 1912: 61; Wolcott, 1947: 72; Knull, 1951: 280, pl. 3, fig. 9; Papp, 1960: 79; Dillon and Dillon, 1972: 274, pl. 28, fig. 10; Ekis, 1975: 17; Barr, 1975: 12; Gosling, 1980: 70; Peck and Thomas, 1998: 86; Mawdsley, 1999: 43, 2002b: 21; Opitz, 2002: figs. 51, 53.

Phyllobaenus pallipennis (Say), 1825 (Figs. 5-5, 5-6)

= Phyllobaenus quadrimaculatus Dejean, 1833 = Hydnocera serrata Newman, 1838

Phyllobaenus pallipennis is recognized by its orangish antennae, mouthparts and legs, dark colored body, pronotum and head, and two yellowish markings on each elytron. The elytral maculations vary in size from one specimen to another, ranging from distinctly separate maculae to contiguous swellings. The margins of the apical third of the elytra are strongly serrulate, with pointed serrulations whose length is greater than the diameter of nearby punctures. The surface of the apical third of each elytron has an anteapical depression. Adults are 3.5 to 5 mm in length.

According to current literature, P. pallipennis may resemble some variations of the likewise variable P. verticalis. Phyllobaenus pallipennis appears to have four distinct, although

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occasionally adjacent, elytral maculae and there are no yellowish markings on its pronotum or head. (See the discussion of Phyllobaenus). Wolcott (1909: 100) observed much variation in the color markings of this species.

Biology: In the summer months of June and July P. pallipennis has been collected in significant numbers on gray birch feeding on aphids. During these months, P. pallipennis prey on many soft-bodied insects, but none more so than aphids. The clerid was rarely collected from neighboring tree species (Chapin, 1948: 304). This species is also a predator of the cotton boll weevil in Texas (Knull, 1951: 283).

Gosling (1980: 71) collected P. pallipennis from sumac and oak and observed it at ultraviolet lights. It has been collected from Rhus copalina Linnaeus (Anacardiaceae).

Mawdsley (2002b: 21) noted that it breeds in the trunks, logs and limbs of oaks and hickory, but that it may occur on flowers and foliage of herbaceous plants.

In Florida adults have been collected in April and May. Specimens have been collected from Malaise traps and sticky traps on slash pine.

Distribution: Alabama, Colorado, District of Columbia, Florida, Indiana, Illinois, Iowa,

Kansas, Louisiana, Maine, Massachusetts, Michigan, Missouri, Nebraska, New Hampshire, New

Jersey, North Carolina, Ohio, Pennsylvania, New York, South Carolina, Texas, Wisconsin, and

Ontario (Wolcott, 1909: 100, 1947: 73; Papp, 1960: 79; Barr, 1975: 12). Wickham and Wolcott

(1912: 64) recorded it from West Springfield, Massachusetts, which is within 10 miles of

Connecticut, and Sparta, Wisconsin, which is within 30 miles of Minnesota. Knull (1951:283) recorded this species from Ohio in Noble County, which is within 50 miles of West Virginia, and

Cincinnati, which is on the Kentucky border. In Florida it has been recorded from Alachua,

Baker, Liberty, and Marion Counties.

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Specimens examined (7): FLORIDA: Alachua County: Gainesville, SE Kincaid Rd., old

field edge, xeric oak habitat, 6 meter Malaise trap, 8-15-V-1999, B.D. Sutton (1, FSCA); Baker

County: Olustee, sticky traps in slash pine, 20-IV-20-V-1978, R.A. Belmont (1, FSCA); Liberty

County: Torreya State Park, 26-IV-1991, M.C. Thomas, 6-V-1989, 7-V-1989, R. Turnbow, 4-V-

2007, J.M. Leavengood, Jr. (1, FSCA; 2, RHTC; 1, JMLC); Marion County: Summerfield, La

Casta subdivision, Rhus copalina, 16-IV-1994, R.E. Woodruff (1, FSCA).

Selected references: Say, 1825: 176; Dejean, 1833: 113; Newman, 1838: 379, 1840: 363; LeConte, 1849: 28, 1859a: 283; Hamilton, 1895: 335; Lohde, 1900: 93; Drury, 1902: 149; Felt, 1906: 693; Wolcott, 1909: 100; Pierce, Cushmen and Hood, 1912: 41; Wickham and Wolcott, 1912: 64; Wolcott, 1947: 73; Chapin, 1948: 304; Knull, 1951: 283, pl. 3, fig. 12; Papp, 1960: 79; Dillon and Dillon, 1972: 274, pl. 29, fig. 4; Ekis, 1975: 17; Barr, 1975: 12; Gosling, 1980: 71; Peck and Thomas, 1998: 86; Mawdsley, 2002: 21; Opitz, 2002: figs. 39, 44.

Phyllobaenus pubescens (LeConte), 1849 (Figs. 5-7, 5-8)

Phyllobaenus pubescens has a blackish blue-green body which appears duller in color due to its dense pubescence. The legs, mouthparts and antennae are orangish yellow. The margins of the apical third of the elytra are weakly and bluntly serrulate. If any of the serrulations appear somewhat pointed, then the length of the serrulation should not exceed the width of nearby elytral punctures. The apical third of the elytra is convex. Adults are about 4 mm in length.

Phyllobaenus pubescens resembles P. humeralis, but lacks the orange humeral maculations and dark colored legs. Phyllobaenus rufipes is also similar in body and leg color, but P. rufipes lacks the dense pubescence.

Sabrosky (1934: 68) discussed the similarity of the larvae of P. pubescens to Isohydnocera curtipennis and P. verticalis, and explained how to use the work of Bøving and Champlain

(1920) regarding this species. Bøving and Champlain (1920: 623) described the larvae and

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compared them to some related taxa. It was noted that P. pubescens was not actually reared and

was listed as “Hydnocera pubescens LeConte (?)”.

Biology: This species is a predator of the cotton boll weevil (Anthonomus sp.)

(Coleoptera: Curculionidae) in Texas, and also on Microbracon sp. (Hymenoptera: Braconidae)

cocoons (Knull, 1951: 280). Mawdsley (2002b: 19) considered P. pubescens to be the most common and abundant clerid associated with prairie flora in North America. Its larvae, and presumably adults, are predaceous on the larvae of Stibadium spumosum Grote (Lepidoptera:

Noctuidae) and Heleothis zea (Boddie) (Lepidoptera: Noctuidae), pests of sunflowers and cotton, respectively. It has been recorded visiting Amorpha fruticosa Torrey, Melilotus officinalis

(Linnaeus) Pallas, Cassia sp. (Fabaceae), Asclepias latifolia (Torrey) Rafinesque

(Asclepiadaceae), Gaillardia sp., Parthenium hysterophorus Linnaeus, Rudbeckia amplexicaulis

Vahl, Rudbeckia grandiflora (Sweet) Gmlin ex De Candolle, Rudbeckia hirta Linnaeus,

Sideranthus rubiginosus (Torrey and Gray) Britton ex Rydberg, and Vernonia interior Smith,

Helianthus petiolaris Nuttall, Helianthus annuus Linnaeus, Aster tanacetifolius Kunth

(Asteraceae), various cactus flowers, Malvastrum apiculatum (Linnaeus) Gray, Callirhoe involucrata (Torrey and Gray) Torrey (Malvaceae), Polytaenia nuttalli De Candolle, Cicuta maculata Linnaeus (Apiaceae), Croton punctatus Jacquin (Euphorbiaceae), Monarda pectinata

Nuttall, Monarda punctata Linnaeus (Lamiaceae), Opuntia sp. (Cactaceae), and Quamasia sp.

(Liliaceae). It has been reared from the nest of Megachile brevis Say (Hymenoptera:

Megachilidae), the stem of Iva ciliata Willdenow (Asteraceae), the flower head of Rudbeckia

amplexicaulis Vahl (Asteraceae), and insect galls (Sabrosky, 1934: 66; Knull 1951: 280;

Mawdsley, 2002b: 19).

Only one specimen has been collected in Florida. It was in April in an alfalfa field.

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Distribution: Colorado, Florida, Illinois, Kansas, Missouri, Montana, Nebraska, New

Mexico, Ohio, Oklahoma, South Carolina, South Dakota, Texas, and Utah (Wolcott, 1909: 99,

1947: 73; Papp, 1960: 79; Mawdsley, 2002b: 19; Barr, 1975: 12). Wickham and Wolcott (1912:

61) recorded it from Brownsville, Texas, which is on the Mexican border. Knull (1951: 280)

recorded it from Cincinnati, Ohio, which is on the Kentucky border. In Florida a single

specimen from Gadsden County has been collected

Specimens examined (1): FLORIDA: Gadsden County: Quincy, in an alfalfa field, 28-

IV-1970, F.W. Mead (1, FSCA).

Selected references: LeConte, 1849: 28; Lohde, 1900: 94; Wolcott, 1909: 99, 1910c: 856; Wickham and Wolcott, 1912: 61; Pierce, Cushman and Hood, 1912: 41; Bøving and Champlain, 1920: 623; Sabrosky, 1934: 66; Wright and Whitehouse, 1941: 72; Wolcott, 1947: 73; Knull, 1951: 280, pl. 3, fig. 11; Papp, 1960: 79; Barr, 1975: 12; Peck and Thomas, 1998: 86; Mawdsley, 2002b: 19.

Phyllobaenus rufipes (Newman), 1840 (Figs. 5-9, 5-10)

This somewhat metallic blue-green hydnocerine has orangish legs, antennae and

mouthparts. The margins of the apical third of the elytra are moderately to weakly, bluntly

serrulate. The apical third of the elytra is convex. Adults range from 3.5 to 5.5 mm in length.

Phyllobaenus rufipes differs from P. pubescens in that it lacks the luster-muting dense

pubescence. It also lacks the dark legs and humeral maculations of P. humeralis and the dark

legs and pale elytral fasciae of P. unifasciatus.

Biology: In Florida adults have been collected from turkey oak and alfalfa fields. Adults

have been collected March through June, August and September. Specimens have been

collected in Malaise traps, malt bait traps and blacklight traps.

Distribution: Alabama, Arkansas and Florida (Wickham and Wolcott, 1912: 60; Wolcott,

1947: 73; Papp, 1960: 79). Wickham and Wolcott (1912: 60) recorded it from Chunchula,

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Alabama, which is within 20 miles of Mississippi. In Florida it has been recorded from Alachua,

Baker, Escambia, Gadsden, Levy, Marion, Nassau, Okaloosa, Putnam, and Union Counties.

Specimens examined (38): FLORIDA: Alachua County: Gainesville, 5-V-1964, R.E.

White (2, FSCA); Pierce's Homestead, west of Gainesville, malaise trap, 21-III-1976, W.H.

Pierce (1, FSCA); Baker County: Glen Saint Mary, malt bait trap, 18-IV-1960, E.W. Holder, Jr.

(1, FSCA); Escambia County: Pensacola, on turkey oak, 17-V-1960, R.E. Woodruff (6, FSCA);

Gadsden County: Quincy, in an alfalfa field, 28-IV-1970, F.W. Mead (1, FSCA); 11-IV-1960,

H.A. Denmark (1, FSCA); Levy County: 3.8 miles southwest of Archer, malaise trap in rosemary and turkey oak sandhill, 10-IV-1988, 17-IV-1988, 17-23-IV-1988, 23-IV-1988, 15-V-

1988, 21-V-1988, 5-VI-1988, 18-IX-1988, P.E. Skelley (14, FSCA); Marion County: 8 miles southeast of Interlachen, on oak, 21-III-1953, B.K. Dozier (1, FSCA); Nassau County: 1.5 miles east of Boulogne, "powerline cut in turkey oaks", 17-IV-1997, P.E. Skelley (1, FSCA); Fort

Clinch State Park, 26-VIII-1993, L.A. Stange and J. Wiley (1, FSCA); Ralph Simmons State

Forest, 1 mile east of Boulogne, 24-V-2002, P.E. Skelley (3, FSCA); St. Mary's R. State Forest,

WMA 1.5 miles east of Boulogne, 17-IV-1997, P.E. Skelley (1, FSCA); Okaloosa County: 1.5 miles west of Holt, on Bryant Bridge Cut Off Road, at blacklight trap, 14-IV-1989, B. Beck, R.

Woodruff and P. Skelley (2, FSCA); Destin, on turkey oak, 16-V-1960, R.E. Woodruff (2,

FSCA); Union County: Highway 241 at Santa Fe River, 16-IV-1989, C.W. Mills, III (1, FSCA).

Selected references: Newman, 1840: 363; Lohde, 1900: 94; Wickham and Wolcott, 1912: 60; Wolcott, 1947: 73; Papp, 1960: 79; Barr, 1975: 13; Peck and Thomas, 1998: 86.

Phyllobaenus suturalis (Klug), 1842 (Figs. 5-11, 5-12)

= Hydnocera limbata Spinola, 1844

The following discussion is based on limited literature and a small number of specimens identified by an authority. Because of the poor descriptions from dated literature, I question the

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accuracy of such identifications. The antennae, legs, mouthparts and two markings on each

elytron are yellowish. The elytral maculations vary in size from one specimen to another,

ranging from distinctly separate maculae to nearly contiguous swellings. The margins of the

apical third of the elytra are strongly serrulate, with pointed serrulations whose length is greater

than the diameter of nearby punctures. The surface of the apical third of each elytron has an

anteapical depression. Adults are 3.5 to 5 mm in length.

It is doubtful that this species occurs in Florida but revisionary work is required (see the discussion of Phyllobaenus).

Biology: In Florida adults have been collected in Florida in May.

Distribution: Florida, Mexico and possibly Panama (Wolcott, 1927: 4, 1947: 73; Barr,

1975: 13). In Florida it has been recorded from Dixie and Marion Counties. Wickham and

Wolcott (1912: 64) recorded this species from Tampa, Hillsborough County, Florida, collected in

May.

Specimens examined (2): FLORIDA: Dixie County: junction of highways 349 and 351,

30-V-1983, R. Turnbow (1, RHTC); Marion County: 11.6 miles southeast of Belleview, 16-V-

1977, R. Turnbow (1, RHTC).

Selected references: Klug, 1842: 313; Spinola, 1844: 49; Gorham, 1882: 168; Lohde, 1900: 94; Wickham and Wolcott, 1912: 64; Wolcott, 1927: 4, 1947: 73; Ekis, 1975: 18; Barr, 1975: 13; Peck and Thomas, 1998: 86.

Phyllobaenus unifasciatus (Say), 1825 (Figs. 5-13, 5-14)

= Hydnocera punctata Spinola, 1844

This dark bodied and dark legged hydnocerine has a pale transverse fascia at about the middle of the elytra. The antennae, mouthparts and occasionally parts of the legs are yellowish brown. The apical profemora and tibiae, apical meso- and metatibiae, and all tarsi may be

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yellowish. The margins of the apical third of the elytra are moderately serrulate. The apical third of the elytra is convex. Adults are around range 4.5 mm in length.

Phyllobaenus unifasciatus is readily distinguished from P. pubescens and P. rufipes by its lack or orangish legs and dense pubescence. Phyllobaenus unifasciatus lacks orange humeral maculations which separates it from P. humeralis. The single median fascia on each elytron separates it from the other species of Phyllobaenus. Due to the pale fascia and body coloration it is similar to Tarsostenus univittatus. Tarsostenus, however, is a much more elongate-bodied insect with longer (as opposed to “stubby”) antennae.

Biology: Phyllobaenus unifasciatus has been recorded from sumac, walnut, American elm, and white oak infested with Hyperplatys sp. larvae, Psenocerus supernotatus (Say)

(Coleoptera: Cerambycidae), Agrilus defectus Lee (Coleoptera: Buprestidae), Magdalis sp.

(Coleoptera: Curculionidae), and Hylurgopinus rufipes (Eichoff) (Coleoptera: Scolytidae)

(Bøving and Champlain, 1920: 636; Pechuman, 1937: 9; Knull 1934: 34, 1951: 280). Gosling

(1980: 71) has reared it from walnut logs. Mawdsley (2002b: 21) notes that it breeds in the trunks and limbs of oaks and hickory, but that it may occur on flowers and foliage of herbaceous plants. It has also been recorded from insect galls (Sabrosky, 1934: 66). Eliason and Potter

(2000: 555) found larvae in dissected stem galls of Callirhytus cornigera (Osten Sacken)

(Hymenoptera: Cynipidae) on pin oak. It was unsure exactly what the larva predated since many other insects occupied the galls.

A single adult has been collected in Florida in April. In Ohio, Illinois and North Carolina, adults are collected in May, June and July (Wickham and Wolcott, 1912: 60; Knull, 1951: 280).

Distribution: Alabama, Arizona, Colorado, District of Columbia, Indiana, Illinois,

Kansas, Kentucky, Maine, Massachusetts, Michigan, Nebraska, New Jersey, New Mexico, New

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York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Texas, Utah, Ontario

(Wolcott, 1909: 99; Wickham and Wolcott, 1912: 60; Wolcott, 1947: 74; Papp, 1960: 80; Barr,

1975: 14; Gosling, 1980: 71; Eliason and Potter, 2000: 555). In Florida it has only been recorded

from Marion County.

Specimens examined (1): FLORIDA: Marion County: Ocala National Forest, Scrub-SW

salt spring, 10-IV-1948, F.N.Y. Young (1, FSCA).

Selected references: Say, 1825: 176; Spinola, 1844: 54, pl. 40, fig. 4; LeConte, 1849: 27, 1859a: 284; Hamilton, 1895: 335; Lohde, 1900: 95; Drury, 1902: 149; Schaeffer, 1908: 131; Wolcott, 1909: 99, 1910c: 856; Wickham and Wolcott, 1912: 60; Bøving and Champlain, 1920: 636; Knull, 1928: 314, 1932: 43, 1934: 34; Sabrosky, 1934: 66; Pechuman, 1937: 9; Hoffmann, 1942: 7; Wolcott, 1947: 74; Craighead, 1950: 203; Knull, 1951: 279, pl. 2, fig. 6; Papp, 1960: 80; Barr, 1975: 14; Gosling, 1980: 71; Peck and Thomas, 1998; 86; Eliason and Potter, 2000: 555; Mawdsley, 2002b: 21.

Phyllobaenus verticalis Say, 1835 (Figs. 5-15, 5-16)

= P. tenella Say, 1835 = P. brachyptera Klug, 1842 = P. lineaticollis Spinola, 1844

This species of Phyllobaenus has the greatest variation seen in Florida’s fauna in

maculation expression, size, and elongation. Its elytra may yellow only at the humeri to nearly

completely yellow, and its head and pronotum may have both yellow and black maculations. Its

legs, however, are always yellow. The margins of the apical third of the elytra are strongly

serrulate, with pointed serrulations whose length is greater than the diameter of nearby

punctures. The surface of the apical third of each elytron has an anteapical depression. Adults

are 3.5 to 5 mm in length.

Wolcott (1909: 99) observed much variation in this species regarding coloration and head

and thorax structure. The degree of variation implies the possibility that more than one species may be present (see the discussion of Phyllobaenus).

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Sabrosky (1934: 68) discussed the larvae of Isohydnocera curtipennis and P. pubescens, and explained how to use the work of Bøving and Champlain (1920) regarding this species.

Bøving and Champlain (1920: 623) described the larvae and compared them to some related taxa.

Biology: Phyllobaenus verticalis has been recorded from wild grape, hickory, river birch, black oak, linden, and bittersweet infested with Eupogonius pubescens LeConte, Grammoptera exigua (Newman), Phymatodes amoenus (Say) (Coleoptera: Cerambycidae), Agrilus betulae

Fisher, and Agrilus geminatus (Say) (Coleoptera: Buprestidae) (Dillon and Dillon, 1974: 274;

Knull, 1932: 43). Mawdsley (2002b: 21) noted that it breeds in the trunks and limbs of oaks and hickory, but that it may occur on flowers and foliage of herbaceous plants.

Osten Sacken (1861: 68) reared it from cynipid galls on white oak. Eliason and Potter

(2000: 555) found larvae in dissected stem galls of Callirhytus cornigera (Osten Sacken)

(Hymenoptera: Cynipidae) on pin oak. They noted that the beetle develops for up to two years in egg chamber of the gall and likely feeds on the cynipid larva.

In Florida adults have been collected February through October. Specimens have been collected in Lindgren funnel traps baited with ethanol and alpha-beta pinene (separately),

Malaise traps and McPhail traps. It has been collected on oak.

Distribution: Florida, Connecticut, District of Columbia, Illinois, Indiana, Iowa, Kansas,

Kentucky, Massachusetts, Michigan, Minnesota, New Jersey, New Hampshire, New York, North

Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, South Dakota, Texas, Vermont,

Wisconsin, Ontario, and Toronto (Wolcott, 1909: 99; Wickham and Wolcott, 1912: 64; Bøving and Champlain, 1920: 636; Wolcott, 1947: 74; Knull, 1951: 282; Papp, 1960: 80; Barr, 1975: 14;

Mawdsley, 1999: 44; Eliason and Potter, 2000: 555). In Florida it has been recorded from

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Alachua, Dixie, Duval, Gadsden, Leon, Marion, Nassau, Okaloosa, Putnam, Suwannee, Volusia,

and Walton Counties.

Specimens examined (46): FLORIDA: Alachua County: Gainesville, 7-VII-1979, G.B.

Edwards, sweeping oaks, 9-II-1975, H. Greenbaum, V-1953, IX-1961, L.A. Hetrick, 13-VI-

1964, 18-VII-1964, 23-V-1964, R.E. White (8, FSCA); Gainesville, Lindgren funnel with 95%

ETOH bait, 29-III-14-IV-1989, T.H. Atkinson (1, FSCA); Gainesville, Doyle Conner building, in window trap 20 feet above ground, 20-IV-1987, 6-X-1987, P.E. Skelley, malaise trap, 3-IV-

1987, 29-V-1987, 5-VI-1987, P.E. Skelley (5, FSCA); Gainesville, NW 42nd Terr. and 34th Pl.,

Turpentine/frontalin-baited Lindgren funnel trap, VI-2005, J.L. Foltz and J.M. Leavengood, Jr.,

28-V-2001, J.L. Foltz (3, JMLC); Gainesville, SE Kincaid Rd, 1 mile north of Payne's Prairie, edge old field/dry oak hammock, 6 m Malaise trap, 7-14-VI-1998, B.D. Sutton (1, FSCA); Dixie

County: 3.5 miles north of Old Town, 1-V-1978, 17-V-1978, M.C. Thomas (2, FSCA); 3.5 miles north of Old Town, route 349, 5-V-1979, 24-V-1979, M.C. Thomas (2, FSCA); 3.7 miles north of Old Town, 20-V-1978, 18-V-1979, 20-V-1978, 23-VI-1978, R. Turnbow (1, FSCA; 6,

RHTC); Duval County: SE Jacksonville, ethanol-baited Lindgren funnel trap, VI-2007, J.M.

Leavengood, Jr. (1, JMLC); Jacksonville, ethanol-baited Lindgren funnel trap, 7-VI-2007, A.

Silagyi and A. Johnson (2, CAPS); Jacksonville, alpha-beta pinene-baited Lindgren funnel trap,

29-VI-2007, A. Silagyi and A. Johnson (1, CAPS); Gadsden County: Quincy, 21-VIII-1971,

T.M. Neal (1, FSCA); Leon County: Tallahassee, alpha-beta pinene-baited Lindgren funnel trap,

3-X-2006, M. Bentley (2, JMLC); Levy County, at light, 3-IV-1954, H.V. Weems, Jr. (1,

FSCA); Marion County: McIntosh, ethanol-baited Lindgren funnel trap, IV-2007, J.M.

Leavengood, Jr. (2, JMLC); Nassau County: Ralph Simmons State Forest, 1 mile east of

Boulogne, 24-V-2002, P.E. Skelley (1, FSCA); Okaloosa County: 1.7 miles northwest of Holt,

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Florida AandM Research Station, Blackwater River State Forest, 2-IX-1976, L.A. Stange (1,

FSCA); Putnam County: 3 miles east of Melrose, K. Ordway Preserve, 24-III-1999, P.E. Skelley

(1, FSCA); Suwannee County: Suwannee River State Park, malaise trap, 24-VI-14-VII-1977,

J.R. Wiley (1, FSCA); Volusia County: Port Orange, in McPhail trap, 11-VI-1969, J.N. Pott (1,

FSCA); Walton County: 1 mile southwest of Holmes County line on Highway 185, 17-IV-1983,

R. Turnbow (1, FSCA).

Selected references: Say, 1835: 164; Klug, 1842: 313; Spinola, 1844: 51, pl. 40, fig. 1; LeConte, 1849: 29, 1859b: 640; Osten Sacken, 1861: 68; Drury, 1879: 171; Hamilton, 1895: 335; Lohde, 1900: 95; Felt, 1906: 605; Wolcott, 1909: 99, 1910c: 856; Wickham and Wolcott, 1912: 64; Bøving and Champlain, 1920: 623, 636; Knull, 1930: 82; Bøving and Craighead, 1931b: 276; Knull, 1932: 43; Sabrosky, 1934: 68; Beal and Massey, 1945: 76; Wolcott, 1947: 74; Craighead, 1950: 203; Knull, 1951: 282, pl. 2, fig. 7; Papp, 1960: 80; Dillon and Dillon, 1972: 274, pl. 28, fig. 12; Ekis, 1975: 18; Barr, 1975: 14; Gosling, 1980: 71; Peck and Thomas, 1998: 86; Mawdsley, 1999: 43; Eliason and Potter, 2000: 555; Mawdsley, 2002b: 21.

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Figure 5-1. Isohydnocera curtipennis

Figure 5-2. Distribution map of Isohydnocera curtipennis

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Figure 5-3. Phyllobaenus humeralis

Figure 5-4. Distribution map of Phyllobaenus humeralis

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Figure 5-5. Phyllobaenus pallipennis

Figure 5-6. Distribution map of Phyllobaenus pallipennis

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Figure 5-7. Phyllobaenus pubescens

Figure 5-8. Distribution map of Phyllobaenus pubescens

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Figure 5-9. Phyllobaenus rufipes

Figure 5-10. Distribution map of Phyllobaenus rufipes

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Figure 5-11. Phyllobaenus suturalis

Figure 5-12. Distribution map of Phyllobaenus suturalis

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Figure 5-13. Phyllobaenus unifasciatus

Figure 5-14. Distribution map of Phyllobaenus unifasciatus

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Figure 5-15. Phyllobaenus verticalis

Figure 5-16. Distribution map of Phyllobaenus verticalis

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CHAPTER 6 KORYNETINAE

Genus Necrobia Olivier, 1795

Necrobia is represented by three cosmopolitan, economically important species in North

America (Barr, 1962: 127; Papp, 1959: 162). There are six other species from Europe, Argentina and South Africa (Papp, 1959: 165). The three North American species are frequently encountered on dry carrion (Wickham and Wolcott, 1912: 67).

Necrobia is best distinguished by its apical maxillary palpomere which is oval and apically truncate, the broad, short and basally dentate tarsal claws, the ninth and tenth antennomeres distinctly narrowly transverse, entire and coarsely granulate eyes, and the eleven-segmented antennae with a small, compact three-segmented club. Their bodies are mostly to entirely metallic bluish-black to green. The elytral humeri and/or legs are orangish in some species.

Bøving and Champlain (1920: 597) described the larvae of Necrobia.

Selected references: Olivier, 1795: 5; LeConte, 1849: 33; Mulsant and Rey, 1863: 122; LeConte and Horn, 1883: 220; Wickham, 1895: 248; Wolcott, 1910c: 861; Wickham and Wolcott, 1912: 67; Bøving and Champlain, 1920: 597; Chapin, 1924: 279; Bradley, 1930: 107; Bøving and Craighead, 1931a: 56; Wolcott, 1947: 88; Knull, 1951: 311, pl. 12, fig. 47, pl. 13, figs. 50, 51; Papp, 1959: 162; Barr, 1962: 127; Dillon and Dillon, 1972: 280; Opitz, 2002: 280.

Key to the Florida Species of Necrobia

1. Body unicolorous, metallic blue or green ...... N. violacea (Fig. 6-5)

1’. Body not unicolorous, with some parts red to reddish-brown ...... 2

2 (1). Legs and basal antennomeres reddish-brown, all else metallic blue or metallic green ...... N. rufipes (Fig. 6-3)

2’. Legs, prothorax and basal ¼ of elytra reddish-brown, front of head and apical ¾ of elytra metallic blue ...... N. ruficollis (Fig. 6-1)

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Necrobia ruficollis Fabricius, 1775 (Figs. 6-1, 6-2)

Necrobia ruficollis is metallic blue, with the ventral surface of head, entire prothorax, basal elytra, meso- and metasternum and legs reddish-brown. The antennae and abdomen are dark brown. Adults are typically around 6 mm in length.

Biology: Necrobia ruficollis is known as the red-shouldered ham beetle (Peck and Thomas

1998: 86). It is often found on the skin of dead animals and, taking advantage of its fellow carrion feeders, it feeds on the larvae of carrion flies, making it both saprophagous and predaceous in its preferred feeding environments (Knull, 1951: 312; Papp, 1959: 164). It is also a pest of dry fish (Papp 1959: 164).

In Florida it has been intercepted in two separate shipments in bone meal.

Distribution: Alabama, California, Florida, Illinois, New Jersey, Ohio, Tennessee,

Wisconsin, Argentina, England and Israel (Wolcott, 1909: 102, 1947: 88; Wickham and Wolcott,

1912: 67; Knull, 1951: 312; Papp, 1960: 87; Peck and Thomas, 1998: 86; Gerstmeier, 1999: 33).

In Florida it has only been recorded from Hillsborough County.

Specimens examined (12): FLORIDA: Hillsborough County: Tampa, in shipment from

Argentina, in bone meal, 19-VI-1928, U.C. Reluff (5, FSCA); Tampa, in bone meal, 23-V-1923,

P. Thomas (7, FSCA).

Selected references: Fabricius, 1775: 57; Latrielle, 1804: 157; LeConte, 1849: 34; Riley, 1874: 101; Hamilton, 1895: 335; Wickham, 1895: 253; Lohde, 1900: 116; Drury, 1902: 149; Wolcott, 1909: 102, 1910c: 861; Wickham and Wolcott, 1912: 67; Scott, 1919: 101; Chapin, 1924: 280; Corporaal, 1937: 17; Wolcott, 1947: 88; Knull, 1951: 312, pl. 13, fig. 51; Papp, 1959: 164, pl. 47, fig. C, 1960: 87; Dillon and Dillon, 1972: 280, pl. 29, fig. 6; Barr, 1975: 18; Gosling, 1980: 74; Peck and Thomas, 1998: 86; Gerstmeier, 1999: 33; Solervicens, 2002: 92; Kolibac, 2003: 91, fig. 34.

Necrobia rufipes Degeer, 1775 (Figs. 6-3, 6-4)

= Tenebrio dermestoides Piller and Mitterpacher, 1783 = Corynetes glabra Champollion, 1814 = Necrobia amethystinae Stephens, 1832

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= Necrobia mumiarum Hope (in Pettigrew), 1834 = Necrobia pilifera Reitter, 1894 = Necrobia foveicollis Schenkling, 1900 = Necrobia pilifera var. aeneipennis Csiki, 1900 = Necrobia pilifera var. cupreonitens Lauffer, 1905

Necrobia rufipes has historically been placed in three different families (Cleridae,

Dermestidae and Tenebrionidae) due to its unusual appearance and behavior. This metallic blue or green, oval-bodied beetle has orange legs and basal antennomeres one through five. The remaining antennomeres, eyes and venter are blackish brown. The first antennomere is thick and slightly bent. Adults range from 3.5 to 7.0 mm in length.

Biology: Necrobia rufipes is best known as the red-legged ham beetle, has also been called the ham beetle by the USDA in the early 1900’s. Those in the meat industry called it the paper worm. In the Pacific Islands it is known as the copra bug. By any name, N. rufipes is associated with rotting meats, fish and cheese, cured ham and bacon, mammal tissue, guano, bone meal, coconut, garlic, grains, silk, cotton, rattan and salt. However, it does not feed on many of these materials. It instead feeds on other pests that infest products or decaying animal matter (Mallis, 1997: 680; Peck and Thomas, 1998: 86). It is a known predator of the cheese skipper, Piophila casei (Linnaeus) (Diptera: Piophilidae), and blowfly and dermestid larvae

(Simmons and Ellington, 1925: 845). It also displays cannibalistic behavior in the absence of other sources of nutrients. A live specimen of N. rufipes was once collected in the skull of a mummy in Egypt (Knull, 1951: 313; Papp, 1959: 164).

Mallis (1997: 680) provided an overview to the life history, development and control techniques of N. rufipes. Simmons and Ellington (1925: 846) offered a more exhaustive life history and many observations of behavior in adult and larval stages. Presented were notes on

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the eggs, larva and larval behavior, prepupa, and pupa, and a formal description of the larva.

Bøving and Champlain (1920: 598) also described the larvae.

Adults have been collected in Florida in all months of the year except March, September

and December. Because most of the Florida records are from shipments from other continents,

this does not necessarily indicate a flight period. Barring specimens whose labels indicate that

they came from shipments, adults have been captured in February, and April through August.

There is a January record that reads “Jacksonville, copra”, but this most likely indicates that it

was collected in a shipment of copra. Specimens have been recorded at blacklights, attracted to

human feces, in copra and in shipments of bone.

Distribution: Arizona, California, Colorado, Florida, Georgia, Idaho, Illinois, Indiana,

Iowa, Kansas, Maryland, Michigan, New Mexico, New York, North Carolina, Ohio, Oklahoma,

Oregon, South Dakota, Texas, Washington, Argentina, the Bahamas, Brazil, China, Columbia,

Cuba, Honduras, Israel, Jamaica, the Kingdom of Tonga, Mexico, Panama, the Philippines,

Saudi Arabia and Suriname (Wolcott, 1909: 102, 1912: 77, 1927: 4, 1947: 88; Wickham and

Wolcott, 1912: 67; Barr, 1950a: 517; Knull, 1951: 313; Vaurie, 1952b: 36; Papp, 1960: 87; Peck

and Thomas, 1998: 86; Gerstmeier, 1999: 33). In Florida it has been recorded from Alachua,

Broward, Duvall, Hendry, Highlands, Hillsborough, Indian River, Liberty, Marion, Miami-Dade,

and Monroe Counties.

Specimens examined (126): FLORIDA: Alachua County: Archer, 22-IV-1961, R.E.

Woodruff (1, FSCA); Gainesville, 18-II-1947, H.V. Weems, Jr., in museum, 16-XI-1972, S.

Humphries (2, FSCA); 20-V-1969, M.L. May (1, FSCA); Broward County: Port Everglades, in

shipment from Cuba, 15-V-1941, J.W. McGlamery, 2-XI-1938, R. Hart (3, FSCA); Port

Everglades, in the ship's hold, from the Philippines, 25-X-1955, J.B. Guthrie, Jr. (78, FSCA);

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Duval County: Jacksonville, copra, 24-I-1943, E.D. Barcus (3, FSCA); Hendry County:

Clewiston, Lake Okeechobee, in human feces, 20-V-1958, R.E. Woodruff (1, FSCA); Highlands

County: Archbold Biological Station, 1-IV-1973, L.L. Lampert, Jr. (1, FSCA); Hillsborough

County: Tampa, in shipment of coconuts from Cartegena, Columbia, 30-I-1956, J.F. Kearney (4,

FSCA); Tampa, in shipment from Argentina, in bones, 18-VI-1928, U.C. Reluff (4, FSCA);

Tampa, in shipment from Argentina, 28-VIII-1922, N. Mowry (6, FSCA); Indian River County:

Vero Beach, in suction trap, 10-VI-1964, W.L. Bidlingmayer (1, FSCA); Liberty County: 20-

VIII-1941 (1, FSCA); 10 miles northeast of Torreya State Park, on dead dog, 16-IV-1963, R.E.

Woodruff (2, FSCA); Marion County: Ocala, blacklight trap, 27-VII-1977, M.C. Thomas (1,

FSCA); Miami-Dade County: VI-1936, S. Warter (1, FSCA); Cutler, on carrion, 18-V-1957,

D.R. Paulson (2, FSCA); Miami, in shipment from Jamaica, 4-I-1954, I.R. Hargrove (2, FSCA);

Miami, in shipment of straw jackets from the Bahamas, 22-VI-1957, F.J. Formichella (10,

FSCA); 27-VII-1975, M.C. Thomas (1, FSCA); Monroe County: Stock Island, 12-V-1961, R.E.

Woodruff (1, FSCA).

Selected references: DeGeer, 1775: 165, pl. 15, fig. 4; Fabricius, 1781: 65; Piller and Mitterpacher, 1783, 68, pl. 7, fig. 8; Rossi, 1792: 33; Olivier, 1795: 5; Fabricius, 1801: 286; Latrielle, 1804: 156; Schönherr, 1808: 51; Champollion, 1814: 41; Stephens, 1832: 417; Hope (in Pettigrew), 1834: 54, pl. 5, figs. 1-3; Dejean, 1837: 127; Klug, 1842: 350; Spinola, 1844: 101; LeConte, 1849: 34; Rosenhauer, 1856: 71; Lacordaire, 1857: 491; Kiesenwetter, 1863: 693; Riley, 1874: 96; Horn, 1876a: 199; Gorham, 1878: 159; Drury, 1879: 171; Gorham, 1883: 193; Reitter, 1894: 85; Hamilton, 1895: 335; Wickham, 1895: 253; Howard and Marlatt, 1896: 105; Lugger, 1899: 150; Lohde, 1900: 116; Schenkling, 1900: 20; Csiki, 1900: 124; Schenkling, 1902: 332; Lauffer, 1905: 406; Escher-Kundig, 1907: 242; Wolcott, 1909: 102, 1910c: 861; Wickham and Wolcott, 1912: 67; Wolcott, 1912: 77; Bøving and Champlain, 1920: 598; Chapman, 1921: 53; Corporaal, 1922: 93; Chapin, 1924: 281; Blaisdell, 1925: 324; Simmons and Ellington, 1925: 845; Corbett and Dover, 1927: 239; Wolcott, 1927: 4; Corporaal, 1937: 17; Hardy, 1943: 136; Hinton, 1945: 239, 267; Judd, 1949: 52; Wolcott, 1947: 88; Barr, 1950a: 516; Knull, 1951: 312, pl. 13, fig. 50; Vaurie, 1952b: 36; Papp, 1959: 164, pl. 47, fig. B, 1960: 87; Dillon and Dillon, 1972: 280, pl. 29, fig. 5; Barr, 1975: 18; Gosling, 1980: 74; Mallis, 1997: 680; Peck and Thomas, 1998: 86; Gerstmeier, 1999: 33; Opitz, 2002: figs. 108, 116; Solervicens, 2002: 92; Kolibac, 2003: 91.

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Necrobia violacea (Linnaeus), 1758 (Figs. 6-5, 6-6)

= Necrobia quadra Marsham, 1802 = Necrobia angustata Falderman, 1835 = Necrobia chalybea Sturm, 1837 = Necrobia errans Melsheimer, 1846

This robust, oval, metallic blue-green beetle is the only member of the genus Necrobia which lacks any reddish-brown coloration. Adults range from 3 to 4.5 mm in length.

Biology: Necrobia violacea is common on the skin and bones of dead mammals, fish, reptiles and birds. As Dermestes (Coleoptera: Dermestidae) larvae are often attracted to the same stimuli, N. violacea commonly feeds on them. In fact, Karl Linne (Linnaeus) thought the beast was a dermestid beetle and described it under the name Dermestes violaceus (Knull, 1951:

313; Papp, 1959: 162). El-Mallakh (1978: 178) observed that N. violacea not only feeds on the adults and immatures of Sarcophaga (Diptera: Sarcophagidae), but that its larvae occasionally bore into the fly puparia, devour the immature puparia contents, and then seal and pupate inside of the fly puparia.

It has been collected at light.

Distribution: Arizona, California, Florida, Idaho, Indiana, Maryland, Massachusetts,

Missouri, New Jersey, New Mexico, New York, Ohio, Oregon, Pennsylvania, South Dakota,

Texas, Washington, Wisconsin, British Columbia and Ontario (Wolcott, 1909: 102, 1947: 88;

Wickham and Wolcott, 1912: 67; Knull, 1951: 313; Papp, 1960: 87). In Florida it has been recorded from Highlands, Miami-Dade, and Orange Counties.

Specimens examined (3): FLORIDA: Highlands County: Archbold Biological Station, at ultraviolet light, 6-V-1979, L.L. Lampert, Jr. (1, FSCA); Miami-Dade County: Coral Gables (1,

FSCA); Orange County: Orlando, 12-XII-1935 (1, FSCA).

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Selected references: Linnaeus, 1758, 1: 356; Marsham, 1802: 323; Falderman, 1835: 210; Sturm, 1837: 43, pl. 232; Melsheimer, 1846: 307; LeConte, 1849: 34; Riley, 1874: 101; Drury, 1879: 171; Hamilton, 1895: 335; Wickham, 1895: 253; Lohde, 1900: 117; Wolcott, 1910c: 862; Wickham and Wolcott, 1912: 67; Chapin, 1924: 280; Hinton, 1945: 239, 272, 285; Wolcott, 1947: 88; Knull, 1951: 313, pl. 12, fig. 47; Papp, 1959: 162, pl. 47, fig. A, 1960: 87; Dillon and Dillon, 1972: 281, pl. 29, fig. 7; Barr, 1975: 18; El-Mallakh, 1978: 178; Gosling, 1980: 74; Peck and Thomas, 1998: 86; Mawdsley, 1999: 44; Kolibac, 2003: 91; fig. 35.

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Figure 6-1. Necrobia ruficollis

Figure 6-2. Distribution map of Necrobia ruficollis

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Figure 6-3. Necrobia rufipes

Figure 6-4. Distribution map of Necrobia rufipes

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Figure 6-5. Necrobia violacea

Figure 6-6. Distribution map of Necrobia violacea

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CHAPTER 7 TARSOSTENINAE

Genus Tarsostenus Spinola, 1844

Tarsostenus is represented in North America by the cosmopolitan Tarsostenus univittatus

(Rossi) (Barr 1962: 127). Tarsostenus is distinguished by the somewhat triangular apical

palpomeres and simple tarsal claws. The antennae end in a loose three-segmented club,

distinguishing Tarsostenus from Cymatodera and Callotillus, which lack antennal clubs. Bøving

and Champlain (1920: 612) described the larvae and compared them to some related taxa.

Selected references: Spinola, 1844: 287; LeConte, 1849:17; Horn, 1880: 150; LeConte and Horn, 1883: 218; Gahan, 1910: 59; Bøving and Champlain, 1920: 612; Chapin, 1924: 278; Bradley, 1930: 107; Wolcott, 1947: 87; Opitz, 2002: 279.

Tarsostenus univittatus (Rossi), 1792 (Figs. 7-1, 7-2)

= Opilo fasciatus Curtis, 1832 = Tillus succinctus Chevrolat, 1842 = Dupontiella fasciatellus Spinola, 1844 = Opilo albofasciatus Melsheimer, 1846 = Notoxus moerens Westwood (in White), 1849 = Tillus picipennis Westwood (in White), 1849 = Tarsostenus biguttatus Montrouzier, 1860 = Opilo incertus MacLeay, 1872

Tarsostenus univittatus is recognized by its narrow, elongate, dark brown to black body, and elytra with a transverse white fascia. The pronotal surface is irregularly punctate, but less densely punctate in the central region. The pronotal margins are gradually divergent until suddenly rounded and constricted posteriorly. Adults are about 4.0 mm long. Cymatodera spp. and Callotillus eburneocinctus Wolcott are similar in appearance, but are best separated by generic characters.

Biology: Tarsostenus univittatus has been recorded from ash, hickory, persimmon, white oak, Acacia tortilis (Forssk.) Hayne raddiana (Savi) Brenan (Fabaceae), Carya pecan (Marshall)

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Engl. and Graebn. (Juglandaceae), Cercis siliquastrum Linnaeus (Fabaceae), Eucalyptus maculata Hook (Myrtaceae), Ficus retusa Linnaeus (Moraceae), Schinus terebinthifolius Raddi

(Anacardiaceae), and Triplochiton sclerodendron (Obeche) (Sterculiaceae) on plywood and furniture made from plywood infested with Lyctus africanus Lesne, Lyctus brunneus (Stephens),

Lyctus planicollis LeConte, Lyctus parallelocollis Blackburn, Sinoxylon sp., Xylobiops basilaris

(Say), Trogoxylon impressum (Comolli) (Coleoptera: Bostrichidae) (Bøving and Champlain

1920: 628; Knull, 1951: 311; Gerstmeier, 1999: 32). Both T. univittatus and several prey species have been found in lumber and other dry, seasoned wood (Knull, 1951: 311). Gerstmeier (1999:

32) regarded it as an important predator of Lyctus africanus, which causes severe damage to wood products.

St. George (1924: 49) offered detailed observations on mating, development, and oviposition. Additionally, he illustrated and made comparisons between the eggs and larvae of

T. univittatus and its prey. Bøving and Champlain (1920: 613) described the larvae and compared them to some related taxa.

Donisthorpe (1901: 362) considered it to be a mutillid mimic, but no further exploration has been published since then.

In Florida T. univittatus has also been collected from an ethanol-baited Lindgren funnel trap. It has been collected from Swietenia macrophylla King (Meliaceae) lumber from Honduras and Philippine mahogany.

Distribution: California, Florida, Ohio, Texas, Australia, Argentina, Chile, Ecuador,

Guatemala, Haiti, Israel, Italy, Honduras, Peru, the Philippines and West Africa (Wolcott, 1947:

87; Barr, 1950a: 518; Knull, 1951: 311; Blackwelder, 1957: 1418; Papp, 1960: 87; Peck, 1993:

153; Peck and Thomas, 1998: 86; Gerstmeier, 1999: 32; Solervicens, 2002: 91; Kolibac, 2003:

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50). Barr (1950a: 518) noted that the distribution of the species was cosmopolitan to the extent

that it matched the distribution of Lyctus, on which it is predaceous.

In Florida it has been recorded from Alachua, Hillsborough, Manatee, Miami-Dade,

Orange, and Palm Beach Counties. Most records are from ports or interceptions, indicating that this species may not be established in Florida (Peck and Thomas, 1998: 86). Only two

specimens indicate the species may be established in Florida, but the data are inconclusive. They

are from Delray Beach, Palm Beach County, and Tampa, Hillsborough County.

Specimens examined (18): FLORIDA: Alachua County: Gainesville, in wood drum from

West Africa, 24-III-2003, D. Harris and J. Diaz (2, FSCA); Hillsborough County: Tampa,

ethanol-baited Lindgren funnel trap, 11-X-2006, Gaskill (1, CAPS); Tampa, in shipment from

Manila, Philippines, 12-X-1953, A.S. Mason (3, FSCA); Manatee County: Bradenton, dead, in orchard basket from Guatemala, 9-X-2000, M. Runnals (1, FSCA); Miami-Dade County: Miami, via Jacksonville on trailer load of Philippine mahogany, 4-VIII-1961 (1, FSCA); Orange County:

Winter Park, in Swietenia macrophylla lumber from Honduras, 12-IV-1994, W. Thompson (9,

FSCA); Palm Beach County: Delray Beach, 1-IV-1974, F. Mulligan (1, FSCA).

Selected references: Rossi, 1792: 44; Stephens, 1830: 324; Curtis, 1832: 6, pl. 270; Dejean, 1837: 126; Stephens, 1839: 197; Chevrolat, 1842: 277; Klug, 1842: 321; Spinola, 1844: 172, pl. 8, fig. 5; Melsheimer, 1846: 306; LeConte, 1849: 17; Westwood (in White), 1849: 48, 57; Westwood, 1852: 50; Lacordaire, 1857: 452; Montrouzier, 1860: 260; Desmarest, 1860: 254; MacLeay, 1872: 269; Chevrolat, 1874: 283; Gorham, 1876: 64; LeConte, 1873: 334; Lewis, 1892: 188; Blackburn, 1900: 136; Lohde, 1900: 52; Donisthorpe, 1901: 362; Bøving and Champlain, 1920: 613, 628; Chapin, 1924: 278; St. George, 1924: 49; Bøving and Craighead, 1931b: 276; Corporaal, 1937: 16; Wolcott, 1947: 87; Barr, 1950a: 517; Craighead, 1950: 203; Knull, 1951: 310, pl. 12, fig. 46; Blackwelder, 1957: 1418; Papp, 1960: 87; Barr, 1975: 18; Peck, 1993: 153; Mawdsley, 1994: 118; Peck and Thomas, 1998: 86; Gerstmeier, 1999: 32; Opitz, 2002: 279, figs. 101, 112; Solervicens, 2002: 91; Kolibac, 2003: 50.

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Figure 7-1. Tarsostenus univittatus

Figure 7-2. Distribution map of Tarsostenus univittatus

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CHAPTER 8 THANEROCLERINAE

North America is home to three genera of the Thaneroclerinae, Ababa, Thaneroclerus and

Zenodosus. The two genera that occur in Florida are each represented by one species, Ababa tantilla (LeConte) and Thaneroclerus buquet (Lefebvre), which are also the only species of their respective genera in North America. Both of these species are cosmopolitan and feed on stored product pests.

Members of this subfamily are often confused with other stored product pests in the families Tenebrionidae, Colydiidae and Silvanidae. Adults are characterized by broadly dilated protarsi and nearly entire eyes. Foster (1976b: 76) offered an extensive treatment of the larvae of the subfamily.

The Thaneroclerinae were treated at a family level by Kolibac (1992). Gerstmeier (2000) provided a synopsis of the morphology, historical research, biology, molecular research, zoogeography and systematics of the “Thanerocleridae” and Cleridae. For the purposes of this thesis the Thaneroclerinae are retained as a subfamily.

Key to the Florida Genera and Species of Thaneroclerinae

1. Prothorax strongly constricted basally, creating collar-like modification; pronotal margins entire, lacking any serration; presence of a distinct carinate intermetacoxal process which spans half of the anterior length of the ventrite on which it is found...... Thaneroclerus buquet (Fig. 8-3)

1’. Prothorax broadly rounded basally, lacking a collar-like modification; pronotal margins minutely serrate basally; carinate intermetacoxal process absent...... Ababa tantilla (Fig. 8-1)

Genus Ababa Casey, 1897

Ababa is represented in North America by a single species, Ababa tantilla (LeConte), the distribution of which reaches across the eastern, central and southern United States (Barr, 1962:

125).

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Adults are distinguished by their small size (1.8-2.6 mm), dull dark brown color, and convex pronotum which is rounded behind. Unlike other members of Thaneroclerinae, the pronotal punctations are rounded and the pronotal margins are minutely serrate behind the middle. Neither Thaneroclerus nor Zenodosus share this minute serration. Further separating this genus from Thaneroclerus, Ababa lacks the pronotal collar-like modification although it does have a basally rounded and mildly constricted pronotum. Foster (1976b: 76) provided a generic larval description and addressed the best means for distinguishing immature Ababa from those of Thaneroclerus and Zenodosus.

Selected references: Casey, 1897: 653; Wolcott, 1910b: 396, 1911: 125; Schaeffer, 1917: 133; Bradley, 1930: 107; Corporaal, 1939: 358; Wolcott, 1947: 76; Foster, 1976b: 76; Opitz, 2002: 273.

Ababa tantilla (LeConte), 1865 (Figs. 8-1, 8-2)

= Ababa crinita Casey, 1897

Some characters to distinguish this species are given under the genus treatment. Other distinguishing characters include a densely, finely punctured head, small eyes, a pronotum that is more densely punctured than the head, and elytra that are densely punctured with the basal punctures larger than those of the pronotum and becoming smaller apically. Its body is entirely dull brown to dark brown.

Biology: Ababa tantilla has been encountered in tobacco bales from the Philippines

(Corporaal, 1939: 358). It has been reared from touchwood (Polyporaceae: Fomes igniarius

(Linnaeus)) in Texas (Knull, 1951: 287).

Ababa tantilla has been collected in Florida in the months of March, June and July. It has been caught in Lindgren funnel traps baited with alpha-beta pinene and ethanol. In Alachua

County it has been collected on Ilex vomitoria Aiton (Aquifoliaceae) and reared from the shelf fungus Phellinus sp. (Polyporaceae) inhabited by Ciidae.

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Distribution: Alabama, District of Columbia, Florida, Louisiana, Massachusetts,

Missouri, Texas, Mexico, Brazil, and the Philippines (Wolcott, 1910b: 396, 1947: 76; Corporaal,

1939: 358; Papp, 1960: 80). Knull (1951: 287) noted that it may occur in Ohio. In Florida it has been recorded from Alachua, Duval, Monroe, and Santa Rosa Counties.

Specimens examined (3): FLORIDA: Alachua County: Gainesville, on Ilex vomitoria,

25-III-1964, F.W. Mead (1, FSCA); Duval County: alpha-beta pinene-baited Lindgren funnel trap, 7-VI-2007, A. Silagyi and A. Johnson (1, CAPS); Santa Rosa County: Gulf Breeze, ethanol-baited Lindgren funnel trap, 31-VII-2006, C. Street (1, FSCA).

Selected references: LeConte, 1865: 96; Casey, 1897: 654; Lohde, 1900: 49; Wolcott, 1910b: 396; Corporaal, 1939: 358, pl. 2, fig. 23; Wolcott, 1947: 76; Knull, 1951: 287, pl. 13, fig. 53; Blackwelder, 1957: 1416; Papp, 1960: 80; Barr, 1962: 125, 1975: 1; Foster, 1976b: 76; Peck and Thomas, 1998: 85; Opitz, 2002: 273, figs. 4, 6, 15.

Genus Thaneroclerus Lefebvre, 1838

Thaneroclerus is represented in North America by a single species, Thaneroclerus buquet

(Lefebvre). Thaneroclerus can be separated from the somewhat similar Zenodosus by its closed procoxal cavities, its proportionally longer first visible abdominal segment, and its duller reddish brown color. Thaneroclerus has a distinctly carinate intermetacoxal process which spans half of the anterior length of the metasternum. The prothorax of Thaneroclerus is also constricted posteriorly, creating a subtle collar. This collar and the pronotum lacking any marginal posterior serrations separate it from Ababa.

Foster (1976: 76) offered a larval treatment of all members of the Thaneroclerinae. Bøving and Champlain (1920: 618) described the larvae and compared them to some related taxa as well.

Selected references: Lefebvre, 1838: 13; Desmarest, 1860: 247; Gemminger and Harold, 1869: 1739; Chevrolat, 1880: 31; Horn, 1880: 150; LeConte and Horn, 1883: 218; Gahan, 1910: 62; Bøving and Champlain, 1920: 618; Bradley, 1930: 106; Matsumura, 1935: 234; Corporaal, 1939: 348; Wolcott, 1947: 75; Foster, 1976b: 76; Opitz, 2002: 273.

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Thaneroclerus buquet (Lefebvre), 1835 (Figs. 8-3, 8-4)

Some characters to distinguish T. buquet are given under the genus treatment. Other distinguishing characters include a densely, finely punctate head, a densely, coarsely punctate pronotum, and elytra that are more densely, coarsely punctured than the pronotum. It is entirely dull brown. Adults range from 4.7 to 6.3 mm in length.

Biology: Thaneroclerus buquet is a predator of insects in stored products such as spices, coffee, tea, drugs and tobacco. It has been encountered in coffee beans, tobacco, drugs, rice and

tea imported from China infested with Araecerus fasciculatus DeGeer (Coleoptera: Anthribidae),

Lasioderma serricorne Fabricius (Coleoptera: Anobiidae), Tribolium ferrugineum Fabricius

(Coleoptera: Tenebrionidae), and Sitophilus oryzae (Linnaeus) (Coleoptera: Curculionidae)

(Corporaal, 1939: 350; Knull, 1951: 286).

Distribution: Florida, California, Hawaii, New York, the West Indies, Puerto Rico, and

Cuba, as well as South America (Columbia and Brazil), Africa (Somalia), Europe (Netherlands,

Germany and Switzerland), and Asia (Israel, China, Japan, India, Burma, Sumatra, Java, and the

Philippines) (Corporaal, 1939: 350; Gerstmeier, 1999: 28). Knull (1951: 286) suggested that it

may occur in Ohio. Because the reported prey, A. fasciculatus, also occurs in dry seeds and dry

and fresh Citrus, and all of the other infesting beetle species mentioned above are cosmopolitan,

T. buquet may be found anywhere that such infestations occur.

Corporaal (1939: 350) recorded specimens of T. buquet from Tampa, Quincy, and Key

West (Hillsborough, Gadsden and Monroe Counties, respectively). Although it has been

collected at various Florida localities and intercepted at Florida ports, it is considered unlikely

that the species is established in Florida. However, it is established in Cuba. It was originally

introduced from the Old World (Peck and Thomas, 1998: 85).

Specimens examined: No Florida specimens were available.

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Selected references: Lefebvre, 1835: 577, pl. 16, fig. 4; Westwood, 1838: 13; Klug, 1842: 310; Spinola, 1844: 207; Chevrolat, 1880: 31; Lohde, 1900: 49; Gahan, 1910: 63; Wolcott, 1910b: 363; Jones, 1913: 10, pl. 1, figs. 10, 11, 12, 13; Runner, 1919: 32; Bøving and Champlain, 1920: 621; Chapin, 1924: 253; Bøving and Craighead, 1931b: 276; Matsumura, 1935: 234; Corporaal, 1937: 14, 1939: 349, pl. 9, fig. 1; Wolcott, 1947: 75; Knull, 1951: 286, pl. 5, fig. 18; Blackwelder, 1957: 1416; Papp, 1960: 80; Barr, 1962: 125, 1975: 1; Ekis, 1975: 10; Foster, 1976: 76; Peck and Thomas, 1998: 85; Gerstmeier, 1999: 28; Opitz, 2002: 273, figs. 2, 5, 8, 10, 12, 26, 49.

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Figure 8-1. Ababa tantilla

Figure 8-2. Distribution map of Ababa tantilla

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Figure 8-3. Thaneroclerus buquet

Figure 8-4. Distribution map of Thaneroclerus buquet

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CHAPTER 9 TILLINAE

Key to the Florida Genera of Tillinae

1. Antennae with 10 segments (but with apparently fewer segments due to strongly reduced size of segments III-V in females or III-IX in males of Monophylla) ...... 2

1’. Antennae with 11 segments ...... 3

2 (1). Antennomere X spatulate and longer than antennomeres I-IX combined, up to four times longer than antennomeres I-IX combined in males; widely distributed throughout Florida (including Miami-Dade and Monroe Counties) and North America ...... Monophylla terminata (Figs. 9-17, 9-18, 9-19)

2’. Antennomere X not spatulate and no longer than antennomeres VIII and IX combined; found only in Miami-Dade and Monroe Counties ...... Callotillus eburneocinctus (Fig. 9-1)

3 (1). Antennomere XI elongate, at least longer than the two preceding segments combined ...... Lecontella brunnea (Fig. 9-15)

3’. Antennomere XI shorter than the two preceding segments combined ...... 4

4 (3). Body length 9.0 mm or larger; pronotum variable ...... Cymatodera

4’. Body length 7.0 mm or smaller; pronotum orange with apical and basal margins black ...... Cymatoderella collaris (Fig. 9-13)

Genus Callotillus Wolcott, 1911

Callotillus is represented by two species across the southwestern United States, Florida,

Mexico and Central America (Barr 1962: 124). The genus is easily confused with Enoclerus.

However, Enoclerus has a three-segmented antennal club whereas Callotillus lacks a club. The

apical antennomere of Callotillus is approximately as long as the preceding two segments

(antennomeres 8 and 9).

Selected references: Wolcott, 1911: 115; Barr, 1962: 124; Opitz, 2002: 275.

Callotillus eburneocinctus Wolcott, 1911 (Figs. 9-1, 9-2)

Callotillus eburneocinctus is the only tilline in North America with a swollen, pale,

transverse fascia near the middle of the elytra. While some members of Cymatodera have a

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midelytral transverse fascia, their fasciae oscillate with irregular margins across the elytra and

are not swollen. Additionally, Callotillus proportionately has much shorter antennae and a more

stout body form.

Callotillus eburneocinctus has a reddish-brown head, antennae, pronotum, thoracic

sternites, and basal elytra. Its abdomen and posterior elytra are bluish-black. Separating the

bluish-black and reddish-brown regions of the elytra are midelytral pale transverse fasciae. It also has long black setae on the middle of the pronotum, arranged in one dense tuft of long setae in the basal region of each elytron, and a dense tuft of setae (shorter than those located basally) on the apical region of each elytron. These tufts, most notably those of the basal elytra, are shared in no other clerid taxon occurring Florida.

Biology: Foster and Barr (1972: 123) noted C. eburneocinctus emerged from buttonwood

(Combretaceae: Conocarpus erecta Linnaeus) in March 1971. The wood was collected by R. L.

Westcott at Tavernier, Key Largo, Florida in November 1970. It has also been reared from

poisonwood (or Florida poisontree) (Anacardiaceae: Metopium toxiferum Linnaeus) and has been collected on white mangrove (Combretaceae: Laguncularis racemosa (Linnaeus)).

In Florida it has been collected in May and June, and has been reared from wood from

February through April and July through September.

Distribution: Callotillus eburneocinctus is known only from the southern extremes of

Florida. It has been recorded from Miami-Dade and Monroe Counties. Wolcott (1911: 116) recorded it only from Key West, Monroe County, Florida, the type-locality. Foster and Barr

(1972: 123) recorded it from Key Largo, Florida. It has been collected on at least six of the

Florida Keys, ranging from Key Largo to Key West.

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Specimens examined (28): FLORIDA: Miami-Dade County: Miami, 25-VI-1965, B.K.

Dozier (1, FSCA); Miami, Virginia Key, 13-VI-1963, B.K. Dozier, on Conocarpus erecta L., 23-

VI-1970, G.H. Nelson, on Laguncularis racemosa (L.), 22-VI-1970, G.H. Nelson (5, FSCA);

Monroe County: Big Pine Key, reared from wood, emerged II-1978, emerged IV-1977, emerged

5-IV-1978, emerged 17-IV-1978, emerged 21-IV-1978, emerged 26-IV-1978, emerged 27-IV-

1978, emerged 27-VII-1977, emerged 11-VIII-1977, emerged 20-VIII-1977, emerged 5-IX-

1977, E. Giesbert (13, FSCA); Everglades National Park, Flamingo, 16-V-1991, R. Morris (1,

FSCA); No Name Key, 29-V-1997, R. Turnbow, ex. Metopium toxiferum L., emerged 31-III-

1979, R. Turnbow (3, RHTC); Sugarloaf Key, 2-V-2000, 30-V-1997, R. Turnbow (3, RHTC);

Tavernier, on Key Largo, on Laguncularis racemosa (L.), 19-VI-1970, G.H. Nelson (2, FSCA).

Selected references: Wolcott, 1911: 115; Wickham and Wolcott, 1912: 51; Wolcott, 1947: 67; Papp, 1960: 76; Foster and Barr, 1972: 123; Barr, 1975: 1; Peck and Thomas, 1998: 85.

Genus Cymatodera Gray, 1832

= Cymatoderus Desmarest, 1860

Cymatodera is represented by about 80 species which share a vast distribution, but most

are distributed within the southwestern United States and Northern Mexico (Barr, 1962: 124).

Wickham and Wolcott (1912: 52) considered this genus to be rarely collected and stated that it

would be “unusual to take more than one or two examples from a single plant”. Nearly all

Cymatodera have cryptic color patterns, are nocturnal and are strongly attracted to lights

(Mawdsley, 1994: 117). In daylight, they are more often encountered on stressed or dying

branches than on healthy plant material. Rarely encountered on flowers and occasionally found

resting under bark, Cymatodera is most commonly collected by beating trees and bushes.

Known plant associations include cedars, larch, junipers, acacias, apple, walnut, oaks and

sequoias. Various observations suggest that adults favor the immature stages of cynipoid wasps

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and other gall-inhabiting grubs, fruit tree caterpillars, and the larvae of Scolytidae, Cerambycidae and Buprestidae as prey (Opitz, 2002: 276). Immatures of C. oblita Horn have been collected from the cells of bees and wasps.

Cymatodera is recognized by coarsely granulate eyes, 11-segmented subfiliform antennae with oval, pointed apical antennomeres, and bipartite tarsal claws. Members of Cymatodera range from 4 mm to over 20 mm, but Florida specimens are 7 mm to 20 mm. Lecontella closely resembles Cymatodera (see the discussion of Lecontella for distinguishing characters to separate the two genera).

Bøving and Champlain (1920: 614) described the larvae and compared them to some related taxa. Foster (1976a: 135) offered a description of the larvae of the genus and discusses how to separate them from Monophylla and Lecontella.

Wolcott (1921: 283) reported specimens of an unnamed species from Crescent City,

Florida, and Mississippi which were “similar to C. wolcotti”. He considered them to be a new species but did not describe the species nor did he address the question in future publications.

Selected references: Gray, 1832: 375; LeConte, 1849: 14; Desmarest, 1860: 240; Horn, 1876b: 220; LeConte and Horn, 1883: 217; Wolcott, 1910c: 849; Wickham and Wolcott, 1912: 52; Bøving and Champlain, 1920: 614, 626; Wolcott, 1921: 271; Bradley, 1930: 105; Bøving and Craighead, 1931a: 56; Wolcott, 1947: 68; Barr, 1962: 124, 1972: 1; Foster, 1976a: 135; Opitz, 2002: 276.

Key to the Florida Species of Cymatodera

1. Body bicolorous or tricolorous; elytra with multiple, irregularly margined fasciae ...... 2

1’. Body entirely brown or bicolorous, with a reddish-orange prothorax and the remaining body black to bluish-black; elytra monocolorous or with a pale midelytral fascia ...... 4

2 (1). Elytra tricolorous, the basal third reddish, followed by a yellowish fascia with irregular margins, the apical portion black with yellowish maculae of variable size; only known from Highlands County, Florida ...... C. floridana (Fig. 9-5)

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2’. Elytra bicolorous, basally with at least some evidence of dark brown, followed by five alternating irregularly margined transverse fascia of yellowish and brown to dark brown, with the apex of the elytra brown to dark brown; widely distributed ...... 3

3 (2). Posterior yellow elytral fascia more blotchy than zigzagged, occasionally not reaching elytral margins, sometimes appearing to be two nigh-adjacent maculae which, in combination typically have two or fewer distinct anterior yellow arches, on each elytron; head densely, coarsely punctate; pronotum coarsely punctate, differing in density to the punctation of the basal portion of the head; distributed throughout Florida ...... C. wolcotti (Fig. 9-11)

3’. Posterior yellow elytral fascia consistent in form with those more anteriorly located, sometimes reduced to the swelling of adjacent brown regions, but not appearing blotchy and typically having three or more distinct anterior yellow arches on each elytron; head densely, finely punctate; pronotum densely, finely punctate; distributed farther north, only recorded from Liberty and Putnam Counties in Florida, but is expected to occur throughout Northern Florida ...... C. undulata (Fig. 9-9)

4 (1). Prothorax mostly reddish-yellow (but rarely only on prosternum anterior to the procoxae); elytra black to bluish-black ...... C. bicolor (Fig. 9-3)

4’. Prothorax brown; body and elytra brown ...... C. inornata (Fig. 9-7)

Cymatodera bicolor (Say), 1825 (Figs. 9-3, 9-4)

Cymatodera bicolor is blackish-blue over most of its entire body. Its pronotum is mostly

reddish-orange, nearly always with the exception of a dark band across the anterior and posterior

margins of the pronotum when viewed dorsally. Within the Florida fauna, C. bicolor is quite

obvious, possibly being mistaken only for Cymatoderella collaris, which differs significantly in

size. Cymatodera bicolor is larger measuring about 10 mm in length, about twice the length of

C. collaris. The pronotal disk of C. bicolor has an uneven dorsal surface whereas it is smoothly

convex in C. collaris. Further separating the two, C. collaris has more finely granulate eyes

versus the coarse granulation of C. bicolor.

In C. bicolor mouthparts and two basal-most antennomeres match the orangish color of the

pronotum, which extends to the ventral prothorax. Additionally, the coxae and basal femora

usually match this color. The femora may be orangish in the basal fourth to basal three-fourths.

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However, specimens from northern regions may exhibit no such coloration on the coxae and

femora. The same goes for the coloration of the basal antennomeres. This orangish coloration of

the mouthparts, basal antennomeres, coxae and femora is not shared with C. collaris.

Specimens of C. bicolor from as far north as Michigan and Canada express the reddish-

orange of the prothorax much less. As Wolcott (1921: 278) explains, some specimens from the

northern regions exhibit reddish-orange only on the prosternum anterior to the coxae. Wolcott

(1909: 95) also observed a similar case in some specimens from New York. These northern

specimens had a pronotum that was nearly entirely black, with the exception of the dilated lateral

portions which were described as “pale”. Such specimens may be confused with C. inornata or

Lecontella cancellata, both of which are brown in color. This has been observed in few Florida

specimens. Knull (1951: 276) described C. bicolor as having “a light transverse band at middle of elytron in some specimens”. In Florida specimens with this character seem quite rare and, when present, it is faintly expressed and would likely be overlooked.

Bøving and Champlain (1920: 616) described the larvae and compared them to related taxa.

Biology: This species has been collected from juniper, flowering dogwood, spice bush, pig-nut hickory, birch and sycamore infested with Phloeosinus sp. (Coleoptera: Curculionidae:

Scolytinae), Encyclops coerulea (Say), Microclytus gazellula (Haldeman), Cyrtophorus verrucosus (Olivier) (Coleoptera: Cerambycidae), Ptilinus ruficornis Say and Trichodesma sp.

(Coleoptera: Anobiidae) (Bøving and Champlain, 1920: 626; Knull 1934: 207, 1951: 277).

Surprisingly, Wickham and Wolcott (1912: 53) wrote that C. bicolor is “found occasionally running about in the house but has seldom been” encountered outdoors. Gosling

(1980: 67) observed it at ultraviolet lights in oak-hickory habitats.

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The immatures can be located by following their small, unpacked mines which are tunneled through the sawdust-packed galleries of their prey species. Although they may construct their own pupal cells, they often use the pupal cells of their host. The pupae most often are oriented vertically in the pupal cell.

Adults have been collected in Florida from April to August, and one specimen was captured in December. It has been collected at mercury vapor lights, blacklights, ultraviolet lights, and Lindgren funnel traps baited with ethanol, turpentine and frontalin, and alpha-beta pinene. Of the “specimens examined” listed below, every single specimen of C. bicolor collected from Florida was collected in a trap or at light.

Distribution: Alabama, Arizona, Connecticut, Florida, Georgia, Illinois, Indiana, Iowa,

Kansas, Maine, Maryland, Massachusetts, Michigan, New Jersey, New Mexico, New York,

North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia,

West Virginia, Wisconsin, and Ontario (Wolcott, 1909: 95; Bøving and Champlain, 1920: 626;

Wolcott 1921: 278, 1947: 68; Knull, 1951: 277; Papp, 1960: 77; Barr, 1975: 2; Peck and

Thomas, 1998: 85). It was also recorded from Chicopee, Massachusetts, which is within 50 miles of Rhode Island, and Onaga, Kansas, which is within 50 miles of Nebraska and Missouri

(Wickham and Wolcott, 1912: 53). It has also been recorded from Cincinnati, Ohio, which is on the Kentucky border (Knull, 1951: 276). In Florida it has been recorded from Alachua, Bay,

Escambia, Liberty, Orange, Putnam, and Santa Rosa Counties.

Specimens examined (22): FLORIDA: Alachua County: Gainesville, Beville Hts., blacklight trap, 5-VII-1980, L.A. Stange (1, FSCA); Gainesville, Hogtown Creek and I-75, blacklight trap, 3-VIII-1988, P.E. Skelley (1, FSCA); Gainesville, NW 42nd Terr. and 34th Pl.,

Turpentine/frontalin-baited Lindgren funnel trap, V-2002, J.L. Foltz (1, JMLC); Bay County:,

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alpha-beta pinene-baited Lindgren funnel trap, 1-VI-2007, L. Smith (2, CAPS); ethanol-baited

Lindgren funnel trap, 15-V-2007, L. Smith (1, CAPS); Escambia County: ethanol-baited

Lindgren funnel trap, 4-V-2007, 24-V-2007, 20-VI-2007, C. Street (5, CAPS); Liberty County:

Apalachicola National Forest, ethanol-baited Lindgren funnel trap, V-2007, J.M. Leavengood, Jr.

(1, JMLC); Torreya State Park, ethanol-baited Lindgren funnel trap, V-2007, VI-2007, J.M.

Leavengood, Jr. (5, JMLC); Orange County: University of Central Florida, Orlando, Cypress

Dome, Malaise Trap, 7-VI-1993, S.M. Fullerton (1, UCFC); University of Central Florida,

Orlando, Cypress Forest, Malaise Trap, 26-IV-1999, P. Russell and S.M. Fullerton (1, UCFC);

University of Central Florida, Orlando, Pond Pine Community, ultraviolet light trap, 12-VI-1993,

J.C. Longhurst and S.M. Fullerton (1, UCFC); Putnam County: Lake Winnott, Melrose, ultraviolet light trap, 5-XII-2001, E. Nearns (1, JMLC); Santa Rosa County: 3.5 miles north of

Munson, at light, 10-IV-1999, P.E. Skelley (1, FSCA).

Selected references: Say, 1825: 174; LeConte, 1849: 16, 1859a: 282; Horn, 1876b: 224; Drury, 1879: 171; Hopkins, 1893a: 154, 1893b: 185; Hamilton, 1895: 335; Lohde, 1900: 19; Wolcott, 1909: 95, 1910c: 850; Wickham and Wolcott, 1912: 53; Bøving and Champlain, 1920: 616; Wolcott, 1921: 278; Knull, 1932: 42, 1934: 207; Wolcott, 1947: 68; Craighead, 1950: 199; Knull, 1951: 276, pl. 1, fig. 4; Papp, 1960: 77; Barr, 1975: 2; Gosling, 1980: 67; Peck and Thomas, 1998: 85; Mawdsley, 1999: 40; Opitz, 2002: figs. 9, 11, 21, 27.

Cymatodera floridana Barr, 1972 (Figs. 9-5, 9-6)

This little-collected species has a distinct color pattern which separates it from other

Cymatodera readily. The basal third of the elytra is brown to brownish-orange, an irregularly margined pale yellowish fascia follows, and the apical half of the elytra is black with a single anteapical pale yellowish macula of variable size on each elytron. Each elytral macula may appear to be partially divided, taking the form of two spots which have swollen to contact each other. The color of the head and pronotum is typically a shade darker than that of the basal third of the elytra. The legs and antennae match the color of the pale fasciae and anteapical maculae.

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It head is coarsely, densely punctate with elongate punctures. The prothorax is coarsely, densely punctate. Adults are typically around 9 mm in length.

Two species may be mistaken for C. floridana: C. undulata and C. wolcotti. However both of these similar species have only two different colors on their elytra and their heads and pronota match the color of their elytral humeri.

Biology: In Florida, adults have been collected in January, April, September and

November. Several specimens have been collected from flight-intercept traps.

Distribution: This species is only known from its type locality in Highlands County,

Florida (Barr, 1972: 28). Further literature has not expanded the known distribution.

Specimens examined (7): FLORIDA: Highlands County: Archbold Biological Station, insect flight trap, 21-I-1979, H.V. Weems, Jr. and D.D. Guenther, 8-IV-1878, insect flight trap,

H.V. Weems, Jr. and L.L. Lampert, Jr., insect flight trap, 13-XI-1979, H.V. Weems, Jr. and T.A.

Webber (3, FSCA); Archbold Biological Station, 1977, 11-IV-1978, 24-IX-1978, L.L. Lampert,

Jr., 16-IV-1989, R. Turnbow (3, FSCA; 1, RHTC).

Selected references: Barr, 1972: 27; Barr, 1975: 3; Peck and Thomas, 1998: 85.

Cymatodera inornata (Say), 1835 (Figs. 9-7, 9-8)

= C. tenera LeConte, 1849

Cymatodera inornata is the most plain of the Florida Cymatodera. It is uniformly brown to light brown, and some specimens have a pale, irregularly-margined transverse fascia at about the middle of the elytra. The head is coarsely punctate, the eyes are coarsely granulate, the surface of the pronotum is sparsely, minutely punctate and the elytral surface has coarse punctures arranged in rows, the punctures becoming obsolete apically. Adults are typically around 12 mm in length.

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Cymatodera inornata is similar to Lecontella brunnea (see the discussion of Lecontella for distinguishing characters to separate the two genera). Bøving and Champlain (1920: 617) described the larvae and compared them to some related taxa.

Biology: Cymatodera inornata has been recorded from dry hickory branches infested with

“Bostrichus” bicornis (Web.) (Coleoptera: Bostrichidae), dead standing birch infested with

Melasis (Coleoptera: Eucnemidae) and buprestid larvae, and also from oak, hackberry, beech and redbud (Bøving and Champlain, 1920: 627). A single record of a pupa was found in beech slash infested with anobiid larvae in Delaware County, Ohio (Knull, 1951: 277).

In Florida, adults have been collected from February through July. Specimens have been collected in Lindgren funnel traps baited with turpentine and ethanol, ethanol, and alpha-beta pinene, Malaise traps and blacklight traps.

Distribution: Alabama, Arizona, Arkansas, District of Columbia, Florida, Georgia,

Illinois, Indiana, Iowa, Kansas, Louisiana, Maine, Maryland, Michigan, Missouri, Ohio,

Oklahoma, New Jersey, New York, North Carolina, Pennsylvania, South Carolina, Utah,

Wisconsin, Ontario, Mexico (Wolcott, 1909: 95, 1921: 278, 1947: 69; Papp, 1960: 77; Barr,

1975: 3; Peck and Thomas, 1998: 85). Wickham and Wolcott (1912: 53) recorded it from

Onaga, Kansas, which is within 50 miles of Nebraska. Knull (1951: 277) recorded it from

Cincinnati, Ohio, which is on the Kentucky border. In Florida it has been recorded from

Alachua, Baker, Bay, Escambia, Highlands, Hillsborough, Leon, Liberty, Manatee, and Orange

Counties.

Specimens examined (18): FLORIDA: Alachua County: Gainesville, V-2005, J.M.

Leavengood, Jr., at blacklight trap, 22-III-1979, M.C. Thomas (1, JMLC; 1, FSCA); Baker

County: Olustee, malaise trap, 15-V-1978, R.A. Belmont (1, FSCA); Bay County: alpha-beta

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pinene-baited Lindgren funnel trap, 4-V-2007, L. Smith (1, CAPS); Escambia County: ethanol- baited Lindgren funnel trap, 20-VI-2007, C. Street (1, CAPS); Highlands County: Archbold

Biological Station, at ultraviolet light, 20-IV-1979, L.L. Lampert, Jr. (1, FSCA); Hillsborough

County: Lithia, ethanol-baited Lindgren funnel trap, III-2007, J.M. Leavengood, Jr. (3, JMLC);

Leon County: Tall Timbers Research Station, 20 miles north of Tallahassee, 8-VII-1972, R.L.

Jacques, Jr. (1, FSCA); Liberty County: Apalachicola National Forest, ethanol-baited Lindgren funnel trap, V-2007, J.M. Leavengood, Jr. (1, JMLC); Torreya State Park, ethanol-baited

Lindgren funnel trap, VI-2007, J.M. Leavengood, Jr. (3, JMLC); Manatee County: Myakka

River State Park, FFA/N Plot, Turpentine/ethanol-baited Lindgren funnel trap, 22-III-2001, J.L.

Foltz (1, JMLC); Orange County: Orlando, alpha-beta pinene-baited Lindgren funnel trap, 29-

III-2007, B. Saunders (2, CAPS); Orlando, ethanol-baited Lindgren funnel trap, 16-II-2007, B.

Saunders (1, CAPS).

Selected references: Say, 1835: 161; LeConte, 1849: 14, 15, 1859b: 638; Horn, 1876b: 224; Hopkins, 1893b: 186; Hamilton, 1895: 335; Lohde, 1900: 20; Wolcott, 1909: 95, 1910c: 850; Wickham and Wolcott, 1912: 53; Bøving and Champlain, 1920: 617, 627; Wolcott, 1921: 278; Blackman and Stage, 1924: 43; Bøving and Craighead, 1931b: 276; Wolcott, 1947: 69; Knull, 1951: 277, pl. 1, fig. 1; Papp, 1960: 77; Barr, 1975: 3; Gosling, 1980: 67; Peck and Thomas, 1998: 85.

Cymatodera undulata (Say), 1825 (Figs. 9-9, 9-10)

= C. bosci Chevrolat, 1843 = C. longicollis Spinola, 1844

Cymatodera undulata has a brown head and pronotum. Its elytral humeri, at least somewhat, and apices are brown with zigzagged irregular margins. Immediately following the brown humeri are alternating yellow and brown fasciae with likewise irregular margins. Three of these fasciae are yellowish and there are two brown fasciae within their boundaries. The coloration of the legs matches that of the yellowish fasciae. The head and pronotum are finely, densely punctate. Adults are typically around 10 mm in length.

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The color pattern of Cymatodera undulata is nearly identical to that of C. wolcotti. In C. wolcotti, the punctures are more densely arranged on the head than on the pronotum especially near the attachment of the head. In contrast, the density of the punctures in C. undulata is similar on the head and pronotum. The color pattern of C. floridana is also similar to C. undulata, however C. floridana has three distinct colors on the elytra and its head and pronotum are a darker shade than the basal third of the elytra.

Bøving and Champlain (1920: 616) described the larvae and compared them to some related taxa. Balduf (1926: 29) offered a diagnosis of the pupae.

Biology: Cymatodera undulata has been recorded from chestnut, maple, oak, and Carolina poplar infested with Phymatodes aereus (Newman), Callidium aeneum (DeGeer), Elaphidion sp., Oberea sp. (Coleoptera: Cerambycidae), and Eucnemidae (Coleoptera). It has also been collected on sumac, wild grape, hackberry, birch, butternut, and American bittersweet, and bred from Spanish needles and Virginia creeper (Wickham and Wolcott, 1912: 54; Bøving and

Champlain, 1920: 627; Wolcott, 1921: 282; Knull 1951: 278).

Eliason and Potter (2000: 555) found larvae in dissected stem galls of Callirhytus cornigera (Osten Sacken) (Hymenoptera: Cynipidae) on pin oak. It was unsure exactly what the larva predated since many other insects occupied the galls. Balduf (1926: 29) encountered a female of C. undulata while studying a bur oak gall made by Disholcaspis mamma Walsh

(Hymenoptera: Cynipidae). The larva was found in the pithy substance, just below the outer surface where parasitic wasp larvae tend to burrow, surrounding the central wasp-inhabited core.

The adults and larvae seemed to avoid light exposure when presented the option. It was suggested that the hooks on the ninth abdominal segment are for moving backward by tugging at the top of an inhabited burrow. An immature was observed consuming about 100 chalcidoid

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larvae and an adult was observed consuming, in part or in total, nineteen adult beetles, flies and

true bugs (all of which were phytophagous).

In Florida, adults have been collected in April and May. One specimen was collected at

light.

Distribution: Alabama, Arizona, Arkansas, District of Columbia, Florida, Illinois,

Indiana, Iowa, Kansas, Kentucky, Maryland, Michigan, Missouri, Nebraska, New Jersey, New

York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas,

Virginia, Wisconsin, and Canada (Wolcott, 1909: 96, Wolcott, 1921: 282, 1947: 71; Wickham

and Wolcott, 1912: 54; Papp, 1960: 78; Barr, 1975: 4; Gosling, 1980: 67; Eliason and Potter,

2000: 555). Knull (1951: 278) recorded it in Ohio from Athens, which is within 20 miles of

West Virginia. In Florida it has been recorded from Liberty County. Wickham and Wolcott

(1912: 54) recorded it from Crescent City, Putnam County, Florida, collected in April.

Specimens examined (2): FLORIDA: Liberty County: Torreya State Park, 6-V-1979,

M.C. Thomas, at light, 17-V-1963, H.V. Weems, Jr. (2, FSCA).

Selected references: Say, 1825: 174; Chevrolat, 1843: 31; Spinola, 1844: 147, pl. 10, fig. 1; LeConte, 1849: 15; Horn, 1876b: 228; LeConte, 1859a: 282; Drury, 1879: 171; Hamilton, 1895: 335; Lohde, 1900: 21; Wolcott, 1909: 96, 1910c: 851; Wickham and Wolcott, 1912: 54; Bøving and Champlain, 1920: 616, 627; Wolcott, 1921: 282; Balduf, 1926: 29; Wolcott, 1947: 71; Knull, 1951: 278, pl. 2, fig. 5; Papp, 1960: 78; Ekis, 1975: 16; Barr, 1975: 4; Gosling, 1980: 67; Peck and Thomas, 1998: 85; Eliason and Potter, 2000: 555.

Cymatodera wolcotti Barr, 1950 (Figs. 9-11, 9-12)

= C. confusa Wolcott, 1921

In 1921, Wolcott named this species Cymatodera confusa (Wolcott, 1921: 282; Barr,

1950b: 61), which not to be mistaken for C. modesta var. confusus Spinola. Thus, it was renamed C. wolcotti. Now long understood to be C. wolcotti, this species is often found misidentified in collections due to its strong similarity to C. balteata and C. undulata (Wolcott,

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1921: 283) (see the discussion of C. undulata for distinguishing characters to separate the two

species).

Cymatodera wolcotti has a brown head and pronotum. Its elytral humeri, at least somewhat, and apices are brown with zigzagged irregular margins. Immediately following the brown humeri are alternating yellow and brown fasciae with likewise irregular margins. Three of these fasciae are yellowish and there are two brown fasciae within their boundaries. The coloration of the legs matches that of the yellowish fasciae. The posterior yellow elytral fascia is more “blotchy” than zigzagged, occasionally not reaching elytral margins and sometimes appearing to be two nigh-adjacent maculae which, in combination, typically have two or fewer distinct anterior yellow arches on each elytron. The head is coarsely, somewhat densely punctate. The pronotum is coarsely punctate with variable density, often dense and rarely sparse.

Adults range from 9 mm to 14 mm in length. Foster (1976a: 138) described the larvae.

Biology: In Florida, adults of C. wolcotti have been collected during all months except for

June. Specimens have been collected at mercury vapor and ultraviolet lights, black light traps,

Malaise traps, a wooden block nest intercept trap, Lindgren funnel traps baited with alpha-beta pinene and ethanol (separately) and turpentine and frontalin (together). Adults have been collected on Prunus persica (Linnaeus) Batsch, (Rosaceae) and Quercus nigra Linnaeus

(Fagaceae). They have been reared from fusiform rust canker on Pinus elliottii Engelman

(Pinaceae) and pupae have been collected from cones of the same pine species.

Distribution: Florida, Georgia, North Carolina, Mississippi, Missouri, South Carolina, and

Texas (Wolcott, 1921: 283, 1947: 69; Papp, 1960: 78). Foster (1976a: 138) noted specimens from Harrison County, Mississippi, which is within 50 miles of Alabama and Louisiana. In

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Florida it has been recorded from Alachua, Baker, Clay, Duval, Hillsborough, Jefferson, Liberty,

Miami-Dade, Orange, Pinellas, Putnam, and Suwannee Counties.

Specimens examined (48): FLORIDA: Alachua County: Gainesville, 20-XI-1984, L.R.

Davis, Jr., ex. Prunus persica, 1-V-1978, C. Lieberman, 19-III-1980, G.B. Edwards, 28-XII-

1988, P. Landolt, X-2004, J.M. Leavengood, Jr., IX-1979, 1-X-1983, 8-X-1980, XII-1982, 27-II-

1982, 1-5-III-1974, 2-V-1978, M.C. Thomas (11, FSCA; 1, JMLC); Gainesville, Doyle Conner

Building, blacklight trap, 13-IX-1986, F.W. Mead, malaise trap, 21-I-1973, 5-XI-1973, H.V.

Weems, Jr. (3, FSCA); Gainesville, University of Florida, Surge/Natural Area Drive,

Turpentine/frontalin-baited Lindgren funnel trap in prescribed burn site, III-2000, J.L. Foltz (1,

JMLC); Monteocha, insect flight trap, 6-VII-1977, 3-X-1977, 26-X-1977, J.F. Butler (3, FSCA);

San Felasco Hammock, forest between Sanchez Prairie and hay field, 6m Malaise Trap, 19-IX-

10-X-1997, B.D. Sutton and G. Steck (1, FSCA); Baker County: Osceola National Forest, ex.

Pinus elliottii cones as pupae, emerged 20-X-1956, 18-X-1956, E.P. Merkel (2, FSCA); ex. fusiform rust canker on Pinus elliottii, emerged 6-IX-1961, 24-VII-1961, P. Mikell (1, FSCA);

Clay County: Camp Blanding Training Site, Insect Survey Site 24: Mown Sandhill, N 30o 0.768

W 81o 57.724, light trap, 5-IX-1999, M. Minno and M. Minno (1, FSCA); 2-VII-1960, H.A.

Denmark (1, FSCA); Duval County: Jacksonville, alpha-beta pinene-baited Lindgren funnel trap,

5-I-2007, A. Silagyi (1, CAPS); Hillsborough County: Lithia, ethanol-baited Lindgren funnel trap, III-2007, J.M. Leavengood, Jr. (1, JMLC); Jefferson County: Monticello, 13-VIII-1993 (1,

FSCA); Liberty County: Torreya State Park, 5-V-2007, J.M. Leavengood, Jr., blacklight trap, 17-

IV-1963, E.I. Hazard (1, JMLC; 1, FSCA); Torreya State Park, ethanol-baited Lindgren funnel

trap, II-2007, J.M. Leavengood, Jr. (1, JMLC); Miami-Dade County: Naranja, 10-IX-1962, R.M.

Baranowski (1, FSCA); [no data] (1, FSCA); Orange County: University of Central Florida,

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Orlando, LLP-Sand Pine, Turkey Oak, Malaise Trap, 27-VIII-1997, 7-XI-1997, 9-X-1997, S.M.

Fullerton (3, UCFC); University of Central Florida, Orlando, Sand Pine-Rosemary Scrub,

Malaise Trap, 31-X-1992, 4-X-1992, S.M. Fullerton (2, UCFC); Walt Disney World, S 16 T 24S

R 27E, MW-5 (unburned) sandpine-oak scrub, Malaise Trap, 10-18-X-1996, P. Russell and S.M.

Fullerton (1, UCFC); Wekiwa Springs State Park, burn zone 43/31, S22 T20S R26E, longleaf pine-turkey oak, Malaise Trap, 18-XI-2001, P. Russell and S.M. Fullerton, 28-X-2001, P.

Russell and S.M. Fullerton (2, UCFC); Wekiwa Springs State Park, burn zone LW-5, S39 T19S

R29E, longleaf pine-turkey oak, Malaise Trap, 2-XII-2001, P. Russell and S.M. Fullerton (1,

UCFC); Orlando, alpha-beta pinene-baited Lindgren funnel trap, 30-III-2007, 8-XI-2006, B.

Saunders (3, CAPS; 1, JMLC); Pinellas County: Saint Petersburg, 11-XI-1964, M. Postupack (1,

FSCA); Putnam County: Palatka, at mercury vapor and ultraviolet lights, 10-I-1991, 3-XII-1990,

H.D. Baggett (2, FSCA); Suwannee County: Suwannee River State Park, wooden block nest intercept trap, IV-2004, D. Serrano and J.M. Leavengood, Jr. (1, JMLC).

Selected references: Wolcott, 1921: 282, 1947: 69; Barr, 1950b: 61; Papp, 1960: 78; Barr, 1975: 5; Foster, 1976a: 138, fig. 5; Peck and Thomas, 1998: 85.

Genus Cymatoderella Barr, 1962

Cymatoderella is represented by three species in the southern United States and Central

America (Rifkind, 1993: 280). Species were originally within Tillus, with T. collaris Spinola later becoming the type-species of Cymatoderella. At the time Cymatoderella was described

(Barr, 1962), T. collaris and T. patagoniae Knull were the only species in the genus, a change that eliminated the distribution of the genus Tillus from the New World (Barr, 1962: 123). The genus resembles a small Cymatodera and is best distinguished by its reddish brown pronotum, finely granulate eyes, entire labrum and bifid tarsal claws (see the discussions of Cymatodera for more characters to separate the taxa). Rifkind (1993: 280) emended Barr’s description of the

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genus to accommodate a previously troublesome species, now known as C. morula Rifkind, and offered a key to the North and Central American Cymatoderella.

Selected references: Olivier, 1790, No. 22; Voet, 1806: 78; LeConte and Horn, 1883: 217; Wolcott, 1910c: 849; Chapin, 1924: 174; Bradley, 1930: 105; Bøving and Craighead, 1931a: 56; Wolcott, 1947: 67; Barr, 1962: 123; Rifkind, 1993: 279; Opitz, 2002: 276.

Cymatoderella collaris (Spinola), 1844 (Figs. 9-13, 9-14)

This small tilline has a bluish-black body with a reddish-orange pronotum. The anterior and posterior margins of the pronotum are lined with bluish-black. The head and central pronotum are finely, sparsely punctate. Adults are 4.0 mm to 5.5 mm in length. It may be mistaken for C. bicolor (see the discussion of C. bicolor for characters to distinguish the two species).

Biology: Rifkind (1993: 281) recorded it from Chilean mesquite (Fabaceae: Prosopis chilensis Stuntz) and Acacia sp. (Fabaceae) in Mexico. It has been collected on Quercus nigra

Linnaeus (Fagaceae).

In Florida, adults have been collected in April, May and July. Specimens have been collected from flight-intercept traps and at mercury vapor and blacklights.

Distribution: Alabama, Florida, Georgia, Louisiana, Ohio, South Carolina, Texas, and

Mexico (Wolcott, 1909: 93, 1921: 270, 1947: 68; Papp, 1960: 76; Barr, 1975: 2; Peck and

Thomas, 1998: 85). Wickham and Wolcott (1912: 52) recorded it from Mobile, Alabama, which is within 20 miles of Mississippi. Knull (1951: 275) recorded it from Lake Hope, Ohio, which is within 50 miles of Kentucky and West Virginia. Internationally, C. collaris is reported from

Mexico and Guatemala, but not other Central American countries (Gorham 1886: 332; Wolcott

1909: 94; Rifkind 1993: 280). It is likely that previously published Mexican distributions represent both C. collaris and C. morula Rifkind and records for Guatemala or Honduras are all

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C. morula. In Florida it has been recorded from Alachua, Dixie, Gadsden, Liberty, Orange,

Sumter, and Union Counties.

Specimens examined (23): FLORIDA: Alachua County: Gainesville, 29-IV-1928, G.B.

Merril, on Quercus nigra L., 8-IV-1961, H.V. Weems, Jr., 22-IV-1947, H.V. Weems, Jr., IV-

2003, J.M. Leavengood, Jr. (3, FSCA; 1, JMLC); Highway 241 at the Santa Fe River, 5-IV-1989,

C.W. Mills, III (1, FSCA); San Felasco Hammock State Park, insect flight trap, 2-V-1977, G.B.

Fairchild and H.V. Weems, Jr. (1, FSCA); Dixie County: 3.5 miles north of Old Town, on Route

349, 27-IV-1980, M.C. Thomas (1, FSCA); 4 miles north of Old Town, 6-V-1979, R. Turnbow

(1, FSCA); Gadsden County: Aspalaga Landing, at boat ramp, at mercury vapor and black light,

5-V-2007, J.M. Leavengood, Jr. (1, JMLC); Liberty County: Torreya State Park, 6-V-1989, R.

Turnbow, at light, 16-VII-1987, Matthews and P.E. Skelley, at light, 17-VII-1987, P.E. Skelley,

4-6-V-2007, J.M. Leavengood, Jr. (3, FSCA; 7, JMLC); Orange County: Wekiwa Springs State

Park, burn zone 43/31, S22 T20S R26E, longleaf pine-turkey oak, Malaise Trap, 13-V-2001, P.

Russell and S.M. Fullerton (1, UCFC); Sumter County: I-75 at CR 470, 18-IV-1992, R. Turnbow

(1, FSCA); I-75 at SR 470, 18-IV-1992, M.C. Thomas (1, FSCA); Union County: Highway 241 at the Santa Fe River, 15-IV-1989, C.W. Mills, III (1, FSCA).

Selected references: Spinola, 1844: 98, pl. 2, fig. 6; LeConte, 1849: 12; Gorham, 1886: 332, tab. 13, fig. 7; Lohde, 1900: 11; Drury, 1902: 149; Wolcott, 1909: 93, 1910c: 849; Wickham and Wolcott, 1912: 52; Wolcott, 1921: 270, 1947: 68; Corporaal, 1950: 26; Knull, 1951: 275, pl. 1, fig. 2; Papp, 1960: 76; Barr, 1962: 123, 1975: 2; Ekis, 1975: 16, pl. 3, fig. 12; Rifkind, 1993: 279; Peck and Thomas, 1998: 85; Opitz, 2002: figs. 23, 37.

Genus Lecontella Wolcott and Chapin, 1918

Lecontella is represented by two species in the eastern, central and southwestern United

States and Mexico (Mawdsley, 2002a: 164). Currently, the validity of a third species from southern Florida is under investigation (Opitz and Thomas, personal communication).

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Lecontella resembles Cymatodera, however in Lecontella the apical antennomere is elongate, to a much greater degree in males, and the elytral punctures are very coarse and reach the elytral apices. In Cymatodera, the apical antennomere is not longer than the two preceding segments combined. The antennae of Lecontella are proportionally thicker, looking heavy and less delicate than those of Cymatodera.

Foster (1976a: 134) offered a description of the larvae of the genus based on a single specimen of Lecontella brunnea and discussed how to separate them from Cymatodera and

Monophylla.

Selected references: Wolcott and Chapin, 1918: 107; Bradley, 1930: 105; Wolcott, 1947: 68; Foster, 1976a: 134; Mawdsley, 2002a: 164; Opitz, 2002: 275.

Lecontella brunnea (Spinola), 1844 (Figs. 9-15, 9-16)

= Cymatodera cancellata LeConte, 1854

Older literature lists L. brunnea under Cymatodera. Collection material may include specimens diagnosed as Lecontella cancellata or L. brunnea cancellata. These should be considered L. brunnea. The name L. cancellata was confirmed as a synonym by Ekis (1975: 13).

Previously, Wolcott and Chapin (1918: 107) had designated C. cancellata rather than L. brunnea for the genotype of Lecontella, naming L. brunnea as a homonym.

Lecontella brunnea is easily confused with members of the genus Cymatodera which lack distinct color patterns. It is most similar to C. inornata, which may have a faint, pale, transverse fascia across the middle of the elytra. Without this character, the two are best separated using generic characters. Lecontella gnara Wolcott is the only other member of the genus currently described. It differs most from L. brunnea in that its pronotal punctures are larger and more densely arranged. The eyes of L. brunnea are coarsely granulate and the elytral surface has rows

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of coarse punctures which become smaller approaching the elytral apices. Adults are around 12 mm long.

Biology: This species has been collected from bee nests in dead logs and in the larval cells of the black and yellow mud-dauber, Sceliphron caementarium Drury (Hymenoptera: Sphecidae)

(Knull, 1932; Rau, 1944: 197). Foster and Barr (1972: 123) noted that R. Bittner of Purdue

University reared L. brunnea from the bee species Osmia lignaria Say, Osmia coerulescens

(Linnaeus), and Megachile rotundata (Fabricius) (Hymenoptera: Megachilidae), the wasp species Monobia quadridens (Linnaeus) (Hymenoptera: Vespidae) and Trypargilum striatum

(Provancher) (Hymenoptera: Sphecidae). Bitner (1972: 24) encountered L. brunnea in the nests of Trypargilum clavatum (Say) and Isodontia auripes (Fernald) (Hymenoptera: Sphecidae). The larvae were found in empty sealed and opened cocoons in unbroken nest cells. Adults have also been reared from abandoned Polistes nests (Hymenoptera: Vespidae). It was suspected that a megachilid that reused the abandoned nest cell was parasitized. According to label data,

Mawdsley (2002a: 165) noted that adults were most frequently collected at lights at night.

In Florida, adults have been collected in April through June and September.

Distribution: Arizona, District of Columbia, Florida, Indiana, Iowa, Kansas, Maine,

Maryland, Michigan, Missouri, New Hampshire, New Jersey, North Carolina, Ohio, Oklahoma,

Pennsylvania, Tennessee, and Texas (Wickham and Wolcott, 1912: 53; Wolcott, 1921: 271,

1947: 68; Papp, 1960: 76; Foster and Barr, 1972: 123; Gosling, 1980: 67). Knull (1951: 275) recorded it from Athens, Ohio, which is about 30 miles from the West Virginia border. In

Florida it has been recorded from Alachua, Highlands, Jefferson, Miami-Dade, and Monroe

Counties.

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Currently, William Barr is describing a new species of Lecontella from south Florida

(Thomas, personal communication). As such, it is possible that some specimens examined in this project are not L. brunnea.

Specimens examined (11): FLORIDA: Alachua County: Gainesville, IV-2003, VI-2003,

J.M. Leavengood, Jr., blacklight trap, 18-V-1962, F.W. Mead, V-1967, L.A. Hetrick (2, JMLC;

2, FSCA); Highlands County: Archbold Biological Station, 15-IX-1978, 17-IX-1977, L.L.

Lampert, Jr. (2, FSCA); Archbold Biological Station, 8 miles south of Lake Placid, blacklight trap, 26-V-1007, P.E. Skelley (1, FSCA); Monroe County: Big Pine Key, blacklight, 19-VI-

1973, J.B. Heppner (1, FSCA); Everglades National Park, Flamingo (1, FSCA); Everglades

National Park, Flamingo, 18-V-1975, E. Giesbert (1, FSCA); upper Key Largo, blacklight, 9-V-

1974, R. Turnbow (1, RHTC).

Selected references: Spinola, 1844: 147; Melsheimer, 1846: 306; LeConte, 1849: 15, 1854: 81; Horn, 1876b: 223; Drury, 1879: 171; Wolcott, 1910c: 850; Wickham and Wolcott, 1912: 53; Wolcott and Chapin, 1918: 107; Bøving and Champlain, 1920: 626, Wolcott, 1921: 271; Knull, 1932: 42; Rau, 1944: 197; Wolcott, 1947: 68; Knull, 1951: 275, pl. 2, fig. 8; Papp, 1960: 76; Bitner, 1972: 23; Foster and Barr, 1972: 123; Barr, 1975: 2; Ekis, 1975: 13; Foster, 1976a: 135, figs. 1, 2, 3, 7; Gosling, 1980: 67; Peck and Thomas, 1998: 85; Mawdsley, 2002a: 164.

Genus Monophylla Spinola, 1841

= Macroletus Klug, 1842 = Elasmocerus LeConte, 1849

Monophylla is represented by three North American species across the southwestern, western and eastern states (Opitz, 2002: 274). This sexually dimorphic genus is readily separated from other genera by its elongate, flattened, spatulate apical antennomere. In females, this apical antennomere is often longer than one-third of the length of the entire antenna. In males this antennomere is up to four times the length of all other basal antennomeres. Many members of the genera Chariessa and Neorthopleura possess a similar color scheme to

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Monophylla and also have long antennae with elongate apical antennomeres. However,

Chariessa and Neorthopleura have three modified apical segments which are not straight, while

Monophylla has only one which is rather straight.

The adults and larvae of Monophylla prey primarily on Bostrichidae with cylindrical galleries that are less tightly packed with frass and sawdust than those of Buprestidae. Such galleries are ideal collecting sites for the larvae. Foster (1976a: 134) offered comparative characters to separate the larvae of M. terminata and M. californica from those of Lecontella and

Cymatodera, as well as a detailed description. Bøving and Champlain (1920: 618) also described the larvae and compared them to some related taxa.

Selected references: Spinola, 1841: 75; Klug, 1842: 274; Spinola, 1844: 384; LeConte, 1849: 13; LeConte and Horn, 1883: 217; Hopkins, 1893b: 185; Wolcott, 1910b: 339, 1910c: 849; Schaeffer, 1911: 123; Bøving and Champlain, 1920: 618; Bradley, 1930: 105; Wolcott, 1947: 67; Foster, 1976a: 133; Opitz, 2002: 274.

Monophylla terminata (Say) 1835 (Figs. 9-17, 9-18, 9-19, 9-20)

= M. megatoma Spinola, 1844

This elongate species has a brownish-yellow pronotum with a central black spot which can occupy most of the dorsal pronotum. Its head may be brownish-yellow and it has a brownish- yellow stripe extending from the elytral humerus to the midelytral margin. At the middle of the elytra, there may be pale yellowish macular spurs extending transversely across the elytra from the marginal stripe. This character is similar to the pale, median elytral fascia found in other genera. The size range within the species is great, with males as small as 4.2 mm and females as large as 15.0 mm, however the males and females have similar average lengths (Wolcott, 1909:

93).

Biology: Monophylla terminata is known to feed on bark beetles in conifers and on

various other wood-boring beetles. It has been found in ash lumbar, hackberry, hickory,

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sassafras, wild grape, Celtis sp. (Ulmaceae), American bittersweet (Celastraceae: Celastrus

scandens Linnaeus), and persimmon (Ebenaceae: Diospyros virginiana Linnaeus) infested with

Phymatodes amoenus (Say) (Coleoptera: Cerambycidae), Lyctus sp., Sinoxylon sp. (Coleoptera:

Bostrichidae), Agrilus sp. (Coleoptera: Buprestidae), Pissodes strobi Peck (Coleoptera:

Curculionidae), Scolytus muticus Say, Chramesus sp. (Coleoptera: Curculionidae: Scolytinae), and Anobiidae (Coleoptera) (Hamilton, 1895: 365; Bøving and Champlain, 1920: 626; Knull,

1951: 274). It has been reared from honey locust, mesquite, walnut logs, Vitia sp. (Vitaceae),

Metopium toxiferum Linnaeus (Anacardiaceae), and bamboo (Wickham and Wolcott; 1912: 51;

Wolcott, 1921: 269; Foster, 1976a: 134; Gosling, 1980: 66). One specimen was labeled “ex.

Saururus cernuus”, presumably it was collected from the plant and not reared. This nocturnal species hides beneath bark or in bark crevices during the day.

Mawdsley (1994: 116) considered it to be a part of a tremendous coleopteran mimicry ring with Lampyridae, Cantheridae, Lycidae, Buprestidae, Meloidae, Ptilodactylidae, Elateridae, and the Cleridae Placopterus thoracicus, Pyticeroides laticornis and Chariessa pilosa.

In Florida, adults have been collected from February through September, and have been reared from wood, emerging as late as October under artificial conditions.

Distribution: Alabama, Arizona, District of Columbia, Florida, Georgia, Illinois, Indiana,

Iowa, Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Mississippi, Missouri,

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

Tennessee, Texas, Virginia, West Virginia, Mexico and Ontario (Wolcott, 1909: 93; 1910b: 342,

1921: 269, 1947: 67; Knull, 1951: 274; Papp, 1960: 76; Barr, 1975: 1; Gosling, 1980: 66; Peck and Thomas, 1998: 85). Wickham and Wolcott (1912: 51) recorded it from Onaga, Kansas,

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which is about 30 miles from the Nebraska border, and Iowa City, Iowa, which is within 60

miles of the Wisconsin border.

In Florida it has been recorded from Alachua, Dixie, Hernando, Highlands, Jefferson,

Miami-Dade, and Monroe Counties. Wolcott (1921: 269) recorded it from Crescent City,

Putnam County, Florida.

Specimens examined (33): FLORIDA: Alachua County: Gainesville, 5-V-1968, D.

Habeck, ex. Vitia sp., emerged 24-IX-1990, wood collected 6-IX-1990, M.C. Thomas, 24-IX-

1990, M.C. Thomas (11, FSCA); Dixie County: 3.5 miles north of Old Town, 8-V-1978, M.C.

Thomas (2, FSCA); 3.5 miles north of Old Town, on Route 349, 5-V-1979, M.C. Thomas (1,

FSCA); Hernando County: Withlacoochee State Forest, Richloam Unit, 8-IV-1994, R. Turnbow

(1, RHTC); Highlands County: Archbold Biological Station, 2-V-1978, R. Turnbow, 23-III-

1978, 14-IX-1979, L.L. Lampert, Jr. (1, RHTC; 2, FSCA); Highlands Hammock State Park, 24-

III-1951, H.V. Weems, Jr., ex. Saururus cernuus, 2-III-1950, F.W. Mead (2, FSCA); Jefferson

County: Monticello, 21-V-1994 (1, FSCA); Monroe County: Big Pine Key, emerged V-1977, emerged 2-VIII-1977, emerged 11-VIII-1977, E. Giesbert (3, FSCA); Key Largo, ex. Metopium toxiferum L., emerged 24-VI-1977, emerged 15-VII-1977, emerged 19-VIII-1977, emerged 7-X-

1977, R. Turnbow (7, RHTC); Key Largo, 4-II-1962, B.K. Dozier (1, FSCA); Key Largo, 2

miles north of Tavernier, 20-VI-1973, J.B. Heppner (1, FSCA).

Selected references: Say, 1835: 160; Klug, 1842: 274, pl. 2, fig. 4; Spinola, 1844: 126, 385, pls. 6, 28, figs. 3, 5; LeConte, 1849: 13, 1859b: 637; Gorham, 1876: 61; Drury, 1879: 171; Chittenden, 1890: 154; Hopkins, 1893a: 154; Hamilton, 1895: 335, 365; Lohde, 1900: 9; Fall, 1901: 251; Schenkling, 1903a: 9, pl. 5, fig. 1; Felt, 1906: 501; Wolcott, 1909: 93, 1910b: 342, 1910c: 849; Wickham and Wolcott, 1912: 51; Bøving and Champlain, 1920: 619, 626; Wolcott, 1921: 269; Blackman and Stage, 1924: 43; Champlain and Knull, 1925: 113; Bøving and Craighead, 1931b: 276; Beal and Massey, 1945: 76, 111, 122; Wolcott, 1947: 67; Craighead, 1950: 201; Knull, 1951: 274, pl. 1, fig. 3, pl. 13, fig. 2; Papp, 1960: 76; Ekis, 1975: 12; Foster, 1976a: 134; Barr, 1975: 1; Gosling, 1980: 66; Mawdsley, 1994: 116; Peck and Thomas, 1998: 85; Opitz, 2002: 274, figs. 25, 30, 31.

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Figure 9-1. Callotillus eburneocinctus

Figure 9-2. Distribution map of Callotillus eburneocinctus

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Figure 9-3. Cymatodera bicolor

Figure 9-4. Distribution map of Cymatodera bicolor

178

Figure 9-5. Cymatodera floridana

Figure 9-6. Distribution map of Cymatodera floridana

179

Figure 9-7. Cymatodera inornata

Figure 9-8. Distribution map of Cymatodera inornata

180

Figure 9-9. Cymatodera undulata

Figure 9-10. Distribution map of Cymatodera undulata

181

Figure 9-11. Cymatodera wolcotti

Figure 9-12. Distribution map of Cymatodera wolcotti

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Figure 9-13. Cymatoderella collaris

Figure 9-14. Distribution map of Cymatoderella collaris

183

Figure 9-15. Lecontella brunnea

Figure 9-16. Distribution map of Lecontella brunnea

184

Figure 9-17. Monophylla terminata, female

Figure 9-18. Monophylla terminata, male

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Figure 9-19. Monophylla terminata, male (left) and female (right)

Figure 9-20. Distribution map of Monophylla terminata

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BIOGRAPHICAL SKETCH

John Moeller Leavengood, Jr., was born in New York to John and Leila Leavengood. He was raised in Haddonfield, New Jersey, where he attended Haddonfield Memorial High School and was a member of the tennis and soccer teams until his junior year. John then moved to Saint

Petersburg, Florida and graduated in the class of 2000 of Boca Ciega High School. Initially a business major at the University of Florida, John was eventually swayed to the discipline of entomology by Don Hall and completed his undergraduate studies in December 2003.

Before beginning his graduate degree, John developed an increased interest in beetles while focusing on various projects with James Cuda, John Foltz, and Jorge Pena involving general insect diagnostics. Upon the suggestion of Michael C. Thomas (supervisory committee chair), John pursued the checkered beetles.

During graduate school John worked as a teaching assistant and developed a passion for interacting with students. In the future, John wishes to pursue teaching as a faculty member of a university in Florida so that he may be close to his large family.

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