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Life History and Pesticide Susceptibility of Cybocephalus nipponicus Endrody-Younga (Coleoptera:Cybocephalidae) and a Ta...


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LIFE HISTORY AND PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus ENDRDY-YOUNGA (COLEOPTERA: CYBOCEPHALIDAE) AND A TAXONOMIC REVISION OF THE CYBOCEPHALIDAE OF NORTH AMERICA AND THE WEST INDIES By TREVOR RANDALL SMITH A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2006

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Copyright 2006 by Trevor Randall Smith

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This document is dedicated to Stuart Fulle rton without whom none of this would have been possible. I would also like to dedicate this dissertation to my wife Kathryn who has been very patient throughout my long academic journey.

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ACKNOWLEDGMENTS First and foremost I would like to offer my gratitude to Ronald D. Cave, my advisor. He has helped me in more ways than could ever be listed here. I thank Howard Frank, Michael C. Thomas, Paul E. Skelley, Bill Overholt and Zhenli He for reviews of this manuscript. A special thanks to Bija n Deghan for offering much needed advice and discussion on cycads, reviewing my manuscr ipt and serving on my committee even after his retirement. I also thank Christine Emshousen for tours of and information about the Montgomery Botanical Center. Simon Yu has given endless help and advice concerning experimental design of pesticid e susceptibility studies, and I also extend my gratitude to the curators of numerous collections for loan s of specimens. Jane Medley and Patricia Hope have offered tremendous assistance with figures and photographs and I would like to thank them as well. This research was supported by a grant from the Florida Department of Agriculture and C onsumer Services (DACS 7276186-12). iv

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TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................iv LIST OF TABLES ...........................................................................................................viii LIST OF FIGURES ...........................................................................................................ix ABSTRACT .......................................................................................................................xv CHAPTER 1 LITERATURE REVIEW.............................................................................................1 Cycads ...........................................................................................................................1 General Scale Insect Information .................................................................................2 Cycad Aulacaspis Scale (CAS) ....................................................................................2 Chemical Control of CAS .............................................................................................5 Biological Control of CAS ...........................................................................................6 Cybocephalidae .............................................................................................................6 Cybocephalids as Biol ogical Control Agents ...............................................................7 Rhyzobius lophanthae .................................................................................................11 2 LIFE HISTORY OF Cybocephalus nipponicus ENDRDY-YOUNGA A PREDATOR OF Aulacaspis yasumatsui TAKAGI (HOMOPTERA: DIASPIDIDAE)..........................................................................................................15 Introduction .................................................................................................................15 Materials and Methods ...............................................................................................16 Results and Discussion ...............................................................................................18 3 PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus AND Rhyzobius lophanthae (COLEOPTERA: CYBOCEPH ALIDAE, COCCINELLIDAE)............32 Introduction .................................................................................................................32 Materials and Methods ...............................................................................................34 Insects ..................................................................................................................34 Bioassays Using Coated Glass Vial Method .......................................................35 Statistical Analysis ..............................................................................................36 Results .........................................................................................................................37 v

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Effects of Pesticide on C. nipponicus ..................................................................37 Effects of Pesticide on R. lophanthae ..................................................................37 Discussion ...................................................................................................................37 4 THE CYBOCEPHALIDAE (COLEOPTERA) OF AMERICA NORTH OF MEXICO.....................................................................................................................42 Introduction .................................................................................................................42 Taxonomic History .....................................................................................................45 Materials and Methods ...............................................................................................46 Materials ..............................................................................................................46 Methods ...............................................................................................................47 Definitions ..................................................................................................................48 Cybocephalus Erichson 1844 .....................................................................................49 Key to the species of Cybocephalus of America north of Mexico .............................49 Cybocephalus californicus Horn ................................................................................50 Cybocephalus kathrynae T. R. Smith, New Species ..................................................60 Cybocephalus nigritulus LeConte ..............................................................................63 Cybocephalus nipponicus Endrdy-Younga ..............................................................68 Cybocephalus randalli T. R. Smith, New Species .....................................................73 5 THE CYBOCEPHALIDAE (COLEOPTER A) OF THE WEST INDIES AND TRINIDAD.................................................................................................................91 Introduction .................................................................................................................91 Materials and Methods ...............................................................................................92 Materials ..............................................................................................................92 Methods ...............................................................................................................93 Definitions ..................................................................................................................93 Key to the Cybocephalidae of the West Indies and Trinidad .....................................93 Cybocephalus Erichson 1844 .....................................................................................94 Cybocephalus antilleus T. R. Smith, New Species ....................................................94 Cybocephalus caribaeus T. R. Smith, New Species ..................................................96 Cybocephalus iviei T. R. Smith, New Species ...........................................................99 Cybocephalus nipponicus Endrdy-Younga ............................................................102 Cybocephalus geoffreysmithi T. R. Smith, New Species .........................................103 Pycnocephalus Sharp 1891 .......................................................................................105 Pycnocephalus deyrollei (Reitter), New Combination .............................................106 6 THE CYBOCEPHALIDAE (COLEOPTERA) OF MEXICO.................................119 Introduction ...............................................................................................................119 Material and Methods ...............................................................................................119 Materials ............................................................................................................119 Methods .............................................................................................................120 Definitions ................................................................................................................120 Key to the Cybocephalidae of Mexico.....................................................................121 vi

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Cybocephalus Erichson 1844 ...................................................................................122 Cybocephalus aciculatus Champion.........................................................................122 Cybocephalus New Species 1...................................................................................124 Cybocephalus californicus Horn ..............................................................................126 Cybocephalus New Species 2...................................................................................128 Cybocephalus New Species 3...................................................................................129 Cybocephalus nigritulus LeConte ............................................................................131 Cybocephalus randalli T. R. Smith ..........................................................................132 Cybocephalus schwarzi Champion...........................................................................134 Cybocephalus New Species 4...................................................................................136 Pycnocephalus Sharp 1891 .......................................................................................138 Pycnocephalus metallicus Sharp ..............................................................................138 LIST OF REFERENCES .................................................................................................155 BIOGRAPHICAL SKETCH ...........................................................................................166 vii

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LIST OF TABLES Table page 2-1 Development of Cybocephalus nipponicus on Aulacaspis yasumatsui ...................31 2-2 Mortality (%) of immature stages of Cybocephalus nipponicus during rearing on Aulacaspis yasumatsui .............................................................................................31 3-1 Field rates (1X) for each pesticide used. ..................................................................40 3-2 Percent mortality of Cybocephalus nipponicus per 30 individuals exposed. X = field rate. ...................................................................................................................40 3-3 Percent mortality of Rhyzobius lophanthae per 30 individuals exposed. X = field rate. ...........................................................................................................................40 3-4 Student-Newman-Keuls test showing ranked values of mortality of adult Cybocephalus nipponicus and Rhyzobius lophanthae .............................................41 viii

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LIST OF FIGURES Figure page 2-1 Dorsal habitus. A, Cybocephalus binotatus B, Cybocephalus nipponicus .............25 2-2 Egg of Cybocephalus nipponicus .............................................................................25 2-3 First instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral habitus. .....................................................................................................................26 2-4 Short trumpet-shaped set ae on third instar larva of Cybocephalus nipponicus. ......26 2-5 Third instar larva of Cybocephalus nipponicus A, Dorsal habitus. B, Ventral oblique habitus. ........................................................................................................27 2-6 Pupal chambers of Cybocephalus nipponicus. A, Pupal chamber made from female scale covers. B, Pupal chamber made from sand. ........................................28 2-7 Pupa of Cybocephalus nipponicus A, Dorsal habitus. B, Ventral habitus. .............29 2-8 Adult C. nipponicus, lateral habitus. ........................................................................30 2-9 Larval mortality over time. .......................................................................................30 4-1 Complete Cybocephalus nipponicus genitalia: ED = ejaculatory duct; IS = internal sac; MA = muscle attachment; ML = median lobe; MS = median strut; Tbp = tegmen basal plate; Tdp = tegmen dorsal piece; TS = tegminal strut. ..........77 4-2 Lateral habitus of Cybocephalus randalli................................................................77 4-3 Truncate antenna of C. californicus .........................................................................78 4-4 Emarginate antenna of C. californicus.....................................................................78 4-5 Median lobe, dorsal view, of C. californicus...........................................................78 4-6 Median lobe, dorsal view (slide mounted), of C. californicus .................................78 4-7 Median lobe, lateral view, of C. californicus...........................................................78 4-8 Basal plate, ventral view, of C. californicus............................................................78 ix

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49 U.S. states and Canadian provinces from which specimens of Cybocephalus californicus have been collected. .............................................................................79 4-10 Scutellum of Cybocephaus nipponicus ....................................................................80 4-11 Scutellum of Cybocephalus kathrynae.....................................................................80 4-12 Antenna of C. kathrynae..........................................................................................81 4-13 Median lobe, dorsal view (same as slide mounted), of C. kathrynae .......................81 4-14 Median lobe, lateral view, of C. kathrynae ..............................................................81 4-15 Basal plate, ventral view, of C. kathrynae ...............................................................81 4-16 Collection localities of Cybocephalus kathrynae in Florida. ...................................82 4-17 Antenna of C. nigritulus ...........................................................................................83 4-18 Median lobe, dorsal view, of C. nigritulus...............................................................83 4-19 Median lobe, dorsal view (slide mounted), of C. nigritulus ....................................83 4-20 Median lobe, lateral view, of C. nigritulus..............................................................83 4-21 Basal plate, ventral view, of C. nigritulus ................................................................83 4-22 U.S. states and Canadian provinces from which specimens of Cybocephalus nigritulus have been collected. .................................................................................84 4-23 Antenna of C. nipponicus.........................................................................................85 4-24 Median lobe, dorsal view (same as slide mounted), of C. nipponicus .....................85 4-25 Median lobe, lateral view, of C. nipponicus ............................................................85 4-26 Basal plate, ventral view, of C. nipponicus ..............................................................85 4-27 U.S. states from which specimens of Cybocephalus nipponicus have been collected. ..................................................................................................................86 4-28 Dorsal habitus of C. binotatus ..................................................................................87 4-29 Dorsal habitus of C. nipponicus...............................................................................87 4-30 Antenna of C. randalli.............................................................................................88 4-31 Median lobe, dorsal view, of C. randalli .................................................................88 x

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4-32 Median lobe, dorsal view (slide mounted), of C. randalli.......................................88 4-33 Median lobe, lateral view, of C. randalli .................................................................88 4-34 Basal plate, ventral view, of C. randalli ..................................................................88 4-35 Striations at the apices of elytra on C. randalli........................................................89 4-36 U.S. states from which specimens of Cybocephalus randalli have been collected. ..................................................................................................................90 5-1 Ventral habitus of C. nipponicus............................................................................110 5-2 Ventral habitus of P. deyrollei ...............................................................................110 5-3 Antenna of C. antilleus...........................................................................................111 5-4 Median lobe, dorsal view, of C. antilleus ..............................................................111 5-5 Median lobe, lateral view, of C. antilleus ..............................................................111 5-6 Basal plate, ventral view, of C. antilleus ................................................................111 5-7 Antenna of C. caribaeus .........................................................................................112 5-8 Median lobe, dorsal view, of C. caribaeus............................................................112 5-9 Median lobe, lateral view, of C. caribaeus............................................................112 5-10 Basal plate, ventral view, of C. caribaeus ..............................................................112 5-11 Antenna of C. iviei .................................................................................................113 5-12 Median lobe, dorsal view, of C. iviei .....................................................................113 5-13 Median lobe, lateral view, of C. iviei .....................................................................113 5-14 Basal plate, ventral view, of C. iviei ......................................................................113 5-15 Scutellum of C. iviei ...............................................................................................114 5-16 Scutellum of C. nipponicus....................................................................................114 5-17 Invaginations at the elytral apices of C. iviei .........................................................115 5-18 Antenna of C. nipponicus.......................................................................................116 5-19 Median lobe, dorsal view, of C. nipponicus ...........................................................116 5-20 Median lobe, lateral view, of C. nipponicus ..........................................................116 xi

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5-21 Basal plate, ventral view, of C. nipponicus ............................................................116 5-22 Antenna of C. geoffreysmithi .................................................................................117 5-23 Median lobe, dorsal view, of C. geoffreysmithi .....................................................117 5-24 Median lobe, lateral view, of C. geoffreysmithi.....................................................117 5-25 Basal plate, ventral view, of C. geoffreysmithi ......................................................117 5-26 Antenna of P. deyrollei ..........................................................................................118 5-27 Hind leg of P. deyrollei ..........................................................................................118 5-28 Median lobe, dorsal view, of P. deyrollei ..............................................................118 5-29 Median lobe, lateral view, of P. deyrollei ..............................................................118 5-30 Basal plate, ventral view, of P. deyrollei ...............................................................118 6-1 Antenna of C. aciculatus ........................................................................................142 6-2 Antenna of Cybocephalus new species 1. ..............................................................143 6-3 Median lobe, dorsal view, of Cybocephalus new species 1. ..................................143 6-4 Median lobe, lateral view, of Cybocephalus new species 1. ..................................143 6-5 Basal plate, ventral view, of Cybocephalus new species 1. ...................................143 6-6 Truncate antenna of C. californicus .......................................................................144 6-7 Emarginated antenna of C. californicus .................................................................144 6-8 Median lobe, dorsal view, of C. californicus.........................................................144 6-9 Median lobe, lateral view, of C. californicus.........................................................144 6-10 Basal plate, ventral view, of C. californicus..........................................................144 6-11 Collection localities of Cybocephalus aciculatus, Cybocephalus new species 1 and Cybocephalus californicus in Mexico. ............................................................145 6-12 Antenna of Cybocephalus new species 2. ..............................................................146 6-13 Median lobe, dorsal view, of Cybocephalus new species 2. ..................................146 6-14 Median lobe, lateral view, of Cybocephalus new species 2. ..................................146 6-15 Basal plate, ventral view, of Cybocephalus new species 2. ...................................146 xii

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6-16 Antenna of Cybocephalus new species 3. ..............................................................147 6-17 Median lobe, dorsal view, of Cybocephalus new species 3. ..................................147 6-18 Median lobe, lateral view, of Cybocephalus new species 3. ..................................147 6-19 Basal plate, ventral view, of Cybocephalus new species 3. ...................................147 6-20 Antenna of C. nigritulus .........................................................................................148 6-21 Median lobe, dorsal view, of C. nigritulus.............................................................148 6-22 Median lobe, lateral view, of C. nigritulus............................................................148 6-23 Basal plate, ventral view, of C. nigritulus ..............................................................148 6-24 Collection localities of Cybocephalus new species 2, Cybocephalus new species 3 and Cybocephalus nigritulus in Mexico. .............................................................149 6-25 Antenna of C. randalli...........................................................................................150 6-26 Median lobe, dorsal view, of C. randalli ...............................................................150 6-27 Median lobe, lateral view, of C. randalli ...............................................................150 6-28 Basal plate, ventral view, of C. randalli ................................................................150 6-29 Antenna of C. schwarzi ..........................................................................................151 6-30 Median lobe, dorsal view, of C. schwarzi ..............................................................151 6-31 Median lobe, lateral view, of C. schwarzi ..............................................................151 6-32 Basal plate, ventral view, of C. schwarzi ...............................................................151 6-33 Antenna of Cybocephalus new species 4. 6-33) antenna; 6-34) median lobe, dorsal view; 6-35) median lobe. lateral view; 6-36) basal plate, ventral view. ......152 6-34 Median lobe, dorsal view, of Cybocephalus new species 4. ..................................152 6-35 Median lobe, lateral view, of Cybocephalus new species 4. ..................................152 6-36 Basal plate, ventral view, of Cybocephalus new species 4. ...................................152 6-37 Antenna of P. metallicus ........................................................................................153 6-38 Median lobe, dorsal view, of P. metallicus ............................................................153 6-39 Median lobe, lateral view, of P. metallicus ............................................................153 xiii

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6-40 Basal plate, ventral view, of P. metallicus .............................................................153 6-41 Collection localities of Cybocephalus randalli Cybocephalus schwarzi Cybocephalus new species 4 and Pycnocephalus metallicus in Mexico. ..............154 xiv

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Abstract of Dissertation Pres ented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy LIFE HISTORY AND PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus ENDRDY-YOUNGA (COLEOPTERA: CYBOCEPHALIDAE) AND A TAXONOMIC REVISION OF THE CYBOCEPHALIDAE OF NORTH AMERICA AND THE WEST INDIES By Trevor Randall Smith December 2006 Chair: Ronald D. Cave Major Department: Entomology and Nematology The life history of th e predatory beetle Cybocephalus nipponicus EndrdyYounga was studied by rearing the beetle on the cycad aulacaspis scale, Aulacaspis yasumatsui Takagi. Mean developmental times of egg (7.3 0.8 days), larval (13.7 1.1 days), and pupal (18.6 1.6 days) stages were determined. Mortality in each life stage, adult longevity, and adult sex ra tios also were measured. A clarification of differences between C. nipponicus and C. binotatus Grouvelle is included. The susceptibility of the predatory beetles C. nipponicus Endrdy-Younga and Rhyzobius lophanthae Blaisdell to six pesticides commonly used for treating cycad aulacaspis scale was tested. Three concentrations (half fiel d rate, field rate, and twice field rate) of each pesticide were tested agai nst both beetle species using a coated glass vial bioassay. Nearly 100% mortality in both beetle species occurred at all concentrations when treated with methidathi on, dimethoate, and malath ion. Insecticidal xv

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soap, fish oils, and imidacloprid were much le ss toxic. At half the simulated field rate, C. nipponicus had 33% survivorship with insectic idal soap, 23% survivorship with imidacloprid, and 17% survivorship with fish oil. At half the simulated field rate, R. lophanthae had 56% survivorship with insectic idal soap, 36% survivorship with imidacloprid, and 53% survivorship with fish o il. Mortality rate for each beetle species rose with increasing concentration of each pesticide. The 17 species of Cybocephalidae in Nort h America (including Mexico) and the West Indies are revised. In cluded are redescriptions of Cybocephalus aciculatus Champion, C. californicus Horn, C. nigritulus LeConte, C. nipponicus Endrdy-Younga, C. schwarzi Champion, Pycnocephalus metallicus Sharp and a new combination of P. deyrolli (Reitter). Also included are descriptions of 10 new species: C. antilleus, C. caribaeus C. iviei, C. kathrynae, C. randalli, C. geoffreysmithi and four as yet unnamed new species from Mexico. A key to species illustrations of mo rphological features including detailed drawings of male genitalia, distribu tion data, and host lists are provided. The confusion involving C. nipponicus and C. binotatus Grouvelle is discussed and the differences be tween them are made evident. xvi

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CHAPTER 1 LITERATURE REVIEW Cycads Cycads are an ancient group of plants, so metimes called the coelacanths of the plant world, that date back to the Paleozoic era (Moretti 1990). However, the true rise and dominance of the cycads occurred during th e Mesozoic era. Cycads are believed to be an evolutionary intermediate between ferns and flowering plants (Whiting 1962). Cycads belong to three families: Cycadaceae which includes a single genus, Cycas; Stangeriaceae which also contains only 1 genus, Stangeria ; and Zamiaceae which contains 8 genera, Bowenia, Ceratozamia, Dioon, Encephalartos, Lepidozamia, Macrozamia Microcycas, and Zamia (Whitelock 2002). There are almost 300 species of cycads worldwide, most of which are found in tropical and subtropical environments (Hill 2004). Many of these species are endange red or threatened, with this threat due more to determined collectors than to defo restation, agriculture, a nd urban sprawl (Giddy 1990). While in some cases cycads are used for food or fertilizer, their chief economic importance is as ornamental landscape plants (Thieret 1958). These plants can be found in hundreds of nurseries all over the state of Florida. While older and consequently larger plants can be quite expensiv e, smaller 1 gallon pot size C. revoluta can sell for as little as $10.00 (Home Depot). Aside from its natural beauty and hardiness, Cycas revoluta is a very popular plant with growers because it can be propagated th rough cutting lateral outgrowths or pups. 1

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2 General Scale Insect Information The armored scale insects (Diaspididae) be long to the superfamily Coccoidea in the order Hemiptera. There are over 1,800 desc ribed species of armored scale worldwide (Ben-Dov 1993) and at least 300 occur in North America (Borror et al. 1989). Many of these scales are pests of agriculture and horti culture, generally weakening plants through sap feeding and often causing excessive sooty mold growth on excreted honeydew. In their native habitats scale insects are us ually controlled by natural predators and parasitoids (Hodgson and Martin 2001). The so ft-bodied females and early-instar males of the tribe Diaspidini live beneath a scale covering made of wax secreted by the insect and mixed with shed exuviae of earlier instar s. The scale covering of males is usually smaller and more elongate than that of the fema les. The first instar, or crawler, stage is mobile and able to spread to other plants. Female crawlers will settle and insert their mouthparts into the plant, becoming sessile and remaining in that state throughout their lifetime. Females will molt twice after the cr awler phase, becoming an adult in the third instar. Males become sessile after the crawler phase and remain in that state through the second and third instar before molting into a fourth prepupa stage. The emergent fifth instar male is winged and without mouthparts (Hamon 2000). Cycad Aulacaspis Scale (CAS) At least 20 species of scale insect occur on cycads in Florida, 19 of which cause very little damage to cycads (Dekle 1 976). The cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi, is the most damaging scale found on cycads in Florida (Hodges et al. 2003). This scale is native to Southeast Asia. It has also been found on several islands in the Caribbean, as well as Hawaii (Ben-Dov et al. 2003). In 1992 A. yasumatsui became such a problem in Hong Kong that 70-100% mortality of Cycas

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3 revoluta Thunberg was recorded (Hodgson and Martin 2001) In Guam the endemic Cycas micronesica K.D. Hill has been severely affected by this damaging scale (R. Muniappan 2005, personal commun ication). The first detect ion of the CAS in Florida occurred in 1996 in Miami at the Montgomery Bo tanical Center. The scale is thought to have come in on infested cycads imported from Southeast Asia (Howard and Weissling 1999). By the end of 1997 the scale had spr ead throughout Miami and as far north as Lake Okeechobee and could be found on 20 species of cycads (How ard and Weissling 1999); however it seemed to prefer the genera Cycas and Stangeria a very rare genus. This led to the spread of the scale to Hawaii in 1998 through the legal importation of infested cycads from Florida (Hodgson a nd Martin 2001). Currently CAS has been reported from Pensacola east to Jacksonville and south into the Florida Keys. How far north the scale is actually es tablished is not known. It is suspected that many of the infested cycads in north Florida were in fact transplants from southern nurseries rather than the pests natural progression north. However, this is just speculation. Aulacaspis yasumatsui was first described by Dr. Sadao Takagi from specimens collected in Bangkok, Thailand (Takagi, 1977). This scale will feed on a large number of cycads; however, the most commercially significant are those of the genus Cycas. In Florida, C. revoluta the king sago, and Cycas rumphii Miq., the queen sago, are the most popular cycads used in landscap ing, and unfortunately, both of these cycads are severely attacked by the cycad scale (H oward et al. 1999). Mature A. yasumatsui females have a white armor 1.2-1.6 mm in diameter that is usually of a pyriform shape common in the tribe Diaspidini, with the e xuviae at one end (Ben-Dov 1990). This is by no means the only shape in which this scale may appear. Th e females of this species may have a large

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4 number of shapes and sizes. The male has a very typical, of the tribe Diaspidini, tricarinate, 0.5-0.6 mm-long, white teste with the exuviae at the cephalic end (Howard et al. 1999). In the field the magnolia white scale, Pseudaulacaspis cockerelli (Cooley), often found on cycads, is frequently confused w ith CAS. However, once the covers are removed major differences in the two species can be observed. All st ages of the CAS life cycle, from egg to adult, ar e a uniform orange color, whereas all stages of the magnolia white scale are yellow. Also, the female CAS has a swollen prosoma and is quite compact, whereas the magnolia white scale ha s a slender prosoma and is relatively elongate (Hodges et al. 2003). The magnolia wh ite scale populates more heavily on the adaxial surface, while CAS creates extremel y dense populations on the abaxial surface with relatively few individuals settling on the upper surface (Howard and Weissling 1999). Most importantly, when a cycad is heavily infested with CAS, the sheer volume of individual scales can become so great that the entire plant is coated with a white crust usually made up mostly of male scales (H oward et al. 1996). By contrast, magnolia white scale infestations are much less severe (Howard and Weissli ng 1999). Lastly, CAS infests all parts of the cycad including the leaves, cones, fruits, megasporophylls, stems and roots (Howard and Weissling 1999), whereas the magnolia white scale attacks only the leaves. Before the introduction of CAS the only scale of the genus Aulacaspis in Florida was Aulacaspis rosae (Bouch), a non-native pest of roses (Dekle 1976). Aulacaspis rosae may be a close relative of A. yasumatsui because the 2 nd instar of A. rosae is very

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5 similar morphologically to that of A. yasumatsui (Takagi 1977), but with such different host plants these two scales are not often confused. Chemical Control of CAS There has been some success controlling th e cycad aulacaspis scale with various pesticides. Oils, either an ul tra-fine horticultural oil or a pr oduct containing fish oil, seem to be the most effective chemical control me thod (Hodges et al. 2003). This is not really surprising given that oils have long been used to control ar mored scale insects. The oil not only covers the insects and suffocates them but also covers the surface of the plant making it difficult for crawlers to settle on to the plant (Howard and Weissling 1999). The proper application of the oils is difficult due to the scales tendency to heavily infest the abaxial surface of the leaves, which is difficult to spray (Howard and Weissling 1999). In the case of C. revoluta the architecture of the plant itself, with the margins of the leaflets curling down and inward and fo rming a trough on the abaxial surface of the leaflet, makes foliar oil treatments difficult (Hodges et al. 2003). Frequent (every two weeks) or as needed use of oils seems to be the most effective technique for controlling this scale, and by mixing treatmen ts of oil with treatments of contact insecticides such as malathion or carbaryl, even gr eater scale mortality can be achieved (Hodges et al. 2003). Horticultural oils also seem to at least help control CAS on the root systems of potted cycads. Hodges et al. (2003) found that drench ing the roots of an infested cycad in 2% horticultural oil resulted in 100% mortality of mature females on the roots. However, a root drench would be very difficult to acco mplish properly on field-grown cycads. The use of systemics such as methidathion and dimethoate has yielded mixed results, being very effective in some cases and completely ineffective at control ling the scale in other cases (Hodges et al. 2003). Imidacloprid as a soil drench can be very effective but

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6 Howard and Weissling (1999) found that this product had to be mixed at such high concentrations that not only was the product not labeled for such high rates but also the whole process became extremely uneconomical. They were completely unable to control CAS using imidacloprid at label rates. Ins ect growth regulators such as pyroproxyfen, sold under the trade name Distance, have met with some success controlling CAS (Emshousen and Mannion 2004). Homeowners have found that liberal application of soap can be a very effective control method as well (personal observation, 2004). Biological Control of CAS The cycad aulacaspis scale is considered a pest in Thailand, but native parasitoids are reported to control its populations (Tang et al. 1997). Howard et al. (1999) reported the lack of any native natural enemies in Florida as one of the major reason for the rapid spread of CAS. For this reason two natu ral enemies were imported from Thailand and released by Dr. Richard Baranowski of the UF/IFAS Tropical Research and Education Center at Homestead in 1998. These natural enemies were a parasitic wasp, Coccobius fulvus (Compere and Annecke), and a predacious beetle, which at the time was identified as Cybocephalus binotatus Grouvelle. In Hawaii the coccinellid Rhyzobius lophanthae (Blaisdell) was found to feed quite readily on CAS (Heu and Chun 2000). Howard (1997) mentions the use of R. lophanthae in Florida as a potential predator of CAS. Cybocephalidae The Cybocephalidae differ greatly from the Nitidulidae, the family in which they have been historically placed, not only because they are predatory but also because their basic morphology and anatomy are quite differe nt. Cybocephalid adults have a 4-4-4 tarsal formula instead of 5-5-5 found in Nitidulidae. There are 5 visible ventral plates (leaving out the male anal pl ate) and 5 abdominal spiracle s in cybocephalids instead of

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7 the 6 and 6 that occur in nititdulids. The body of cybocephalids is retractile allowing the mandibles in repose to rest against the metasternum, unlike any other nitidulid. The larvae of Cybocephalidae have a head wit hout dorsal sutures, lack pregomphi and urogomphi on abdominal tergite XI, and ha ve hypostomal rods with divergent hypostomal ridges present posteriorly, hypopha rynx without a sclerome and bracons, maxillae without mola, and annular spiracles w ith 2 lateral air tubes. In contrast, the larvae of Nitidulidae have pregomphi and urogomphi, no hypostomal rods but with hypostomal ridges strongly convergent posteriorly, hypopharynx with a sclerome and bracons, maxillae with raised mola, and biforous spiracles (Kire jtchuk 1997). This has led to the cybocephalids being shuffled around between family (Murray, 1864, Parsons 1943, Endrdy-Younga 1968) and subfamily (Horn 1879, Arnett 1960, Habeck 2002) status. Before the taxonomic study presented here only one species of Cybocephalidae, Cybocephalus nigritulus LeConte, was known to be native to the state of Florida; a second species is described in Chapter 4. Howard and Weissling (1999) reported C. binotatus was released and has subsequently beco me established in south Florida. However, as will be shown later, this species was misidentified and is actually another Asian species, Cybocephalus nipponicus Endrdy-Younga Cybocephalus nipponicus has been widely used in the United States as a biological control agent for the euonymus scale, Unaspis euonymi (Comstock), and for many other pest scale species around the world. Cybocephalids as Biological Control Agents With the exception of coccinellids, species of the genus Cybocephalus are the most important predators of armored scales (Blumberg and Swirski 1982). While

PAGE 24

8 cybocephalids are well-known armored scale feed ers, their effectiven ess as biological control agents have no t been well studied (Drea and Carl son 1988). Even with such a limited understanding of the biology and ecology of this group of beetles, species of Cybocephalus are being released in many areas of the world. There is very little literature on cybocephalids and what there is usually consis ts of a small note about them being released and no evaluation data (Labuschange et al. 1996; Swirski and Wysoki 1995; Hodges et al. 2003). In South Africa, C. nipponicus, along with a parasitoid Aphytis sp., was imported from Thailand and successfully established in commercial mango orchards. In the following four seas ons these beneficial insects seemed to effectively control the mango scale, Aulacaspis tubercularis Newstead, with augmentative releases. A similar release of C. binotatus (but most likely C. nipponicus) to control the Japanese bayberry whitefly, Parabemisia myricae (Kuwana), on citrus and avocado trees in Israel was le ss successful. In this case the researchers were unable to establish a viable population of the predatory beetle in the field (Swirski and Wysoki 1995). Drea and Carlson (1988) and Van Driesche et al. (1998) were able to establish C. nipponicus in the Virginia/Maryland area as well as in New England. In 1995, the New Jersey Department of Agriculture embarked on an aggressive plan to combat the euonymus scale using C. nipponicus as a biological control agent (Hudson et al. 2001). By the year 2000, after many supplemental release and recovery plans had been implemented, it was obvious that the beetles had established populati ons all over the state where they were controlling and in some cases eradicating the euonymus scale.

PAGE 25

9 Others have noted cybocephalids acting as a natural control of diverse species of armored scales, including at least eight spec ies in Turkey (Erler and Tun 2001), coconut scale (Aspidiotus destructor (Signoret)) in Brazil (Lima 2002), and the brown apricot scale (Lecanium corni Bouch) and San Jose scale (Quadraspidiotus perniciosus (Comstock)) (Heintz 2001) in Californ ia. However, in Mauritius the native Cybocephalus mollis Endrdy-Younga seemed unable to control the spread of the sugarcane scale, Aulacaspis tegalensis (Zhnt.), even when in conjunction with three other natural enemies of the scale (Williams and Greathead 1973). Because many scale insects persist year-round, it is important that a biological control agent be found that is also persistent throughout the year. The cybocephalids are uniquely suited for this in that the placemen t of eggs and subsequent development of larvae beneath the armored scale allow them some protection from both the elements and pesticides (Alvarez and Van Driesche 1998a). In Greece, Katsoyannos (1984) found that Cybocephalus fodori Endrdy-Younga was able to surv ive in pesticide-treated fruit orchards. In date palm plan tations in Israel, Kehat et al. (1974) found that while all coccinellids in a chemically trea ted plantation died, species of Cybocephalus survived. The dietary needs of cybocephalids also make them good candidates for use as biological control agents. Alvarez and Va n Driesche (1998a) found that at low scale densities cybocephalids were able to maintain their populations and keep euonymus and San Jose scale populations in check. In the case of C. nipponicus an average of 19.5 scales were attacked over the entire larval lifetime, contra sted with an average of 199 green scales ( Ceroplastes japonicus Green) consumed by the coccinellid Chilocorus kuwanae Silvestri (Xia et al. 1986). Thickness of the scale cover is also a factor in

PAGE 26

10 effectiveness of a beetle as a predator. Blumberg and Swirski (1982) found that C. micans and C. nigreceps were unable to feed on adult female diaspidid scales. It has also been noted that coccinellids are more likely to feed on scales with thin or easily penetrated scale covers (Honda and Luck 1995). Cybocephalus nipponicus seemed to be less affected by scale cover characteristics (A lvarez and Van Driesche 1998a). In fact, C. nipponicus was able to feed on the adult female s of both the San Jose scale and the euonymus scale (Alvarez and Van Drieche 1998 a). In the absence of prey, female cybocephalids are able to withhold eggs for up to 2 days, indicating that oviposition strategy is not only governed by food consumption but by some qualitative features of the scale population (Alvarez and Van Drieche 1998b). In the presence of greater scale densities cybocephalids will in crease their egg production until they reach a constant. Again this allows cybocephalids to maintain their populations at low scale densities. If circumstances allow, female cybocephalid s will lay only one egg under a single scale cover. However, if the number of scales in a patch is low and th e beetle cannot find new scales it will lay eggs under a scale that ha s already been parasiti zed (Alvarez and Van Drieche 1998b). The beetle larvae consume more prey when feeding on young scales; therefore it seems that for greater survivabi lity of their offspring female cybocephalids prefer to oviposit on older scales (Alvar ez and Van Driesche 1998b). Alvarez and Van Driesche (1998b) found that the highest larval survival rate could be found in larvae feeding on scales older than 30 days. It wa s also noted by Blumberg and Swirski (1982) that C. nigriceps very rarely laid eggs under dead female scales. Certain aspects of the biology of Cybocephalus nigriceps (J. Sahlberg), Cybocephalus micans Reitter, and Cybocephalus gibbulus Erichson were studied by

PAGE 27

11 Nohara and Iwata (1988) and Blumberg and Swirski (1982). Tanaka and Inoue (1983) were the first to really st udy the feeding behavior of C. nipponicus Later, Drea and Carlson (1988) and Alvarez and Van Drie sche (1998a, 1998b) cont inued the biology of C. nipponicus. There remains much to be studied about the life cycle, breeding habits, oviposition behavior, or di etary requirements of C. nipponicus to determine whether it has any significant effect as a biological co ntrol agent of the cycad aulacaspis scale. Without some form of baseline data on the be etle, no real effectiven ess research can be done. Rhyzobius lophanthae Rhyzobius (= Lindorus ) lophanthae (Blaisdell) is a sma ll, pubescent coccinellid belonging to the tribe Coccidulini. The a dult is 2.4-2.5 mm in length and between 1.7 and 1.8 mm in width, and has black or brown elytra and an orange-brown thorax and head region. The body form is elongate or ov al with a dense mat of hairs covering the dorsal surface. The head is partly conceal ed beneath the pronotum, with 11-segmented antennae, the last 3 segments of which are broa der then the rest to fo rm a club. The tarsal claws are not toothed. This bee tle is often referred to as th e singular black lady beetle or the scale destroyer. Rhyzobius lophanthae is a coccoidophagous predator native to Australia. It is considered by many to be one of the most economically important natural enemies of armored scale insects (Yus 1973, Rosen 1990, Stathas 2001). This beetle has been released in many areas of the world to contro l a plethora of armore d scale species (Honda and Luck 1995). There are many examples of successful control of scale insects using R. lophanthae especially in the Mediterr anean region (Greathead 1973). There have been some very high profile failures as well, such as the inability of R. lophanthae to control

PAGE 28

12 the California red scale in California (Greathead 1973). Honda and Luck (1995) found that the morphological characteri stics of the scale itself are a major determining factor in how effective R. lophanthae will be in controlling a scale species. They discovered that Rhyzobius mandibles were not as effective as thos e of other species of coccinellids that specialize on armored scales, such as Chilocorus cacti (L.), a species frequently seen on scale-infested cycads in Florida (RD Cave 2006, personal communication). Rhyzobius lophanthae lacks a tooth at the apex used for pryi ng scales from the substrate and the incisor region is neither as acutely angled nor as sharp as that of specialists like C. cacti Being much less specialized and more of a generalist predator has allowed R. lophanthae to be used to control a la rge number of pest species. Rhyzobius lophanthae seem to be especially effective in closed environments such as greenhouses or in botanical gardens (Anonymous 2003). They have been recorded feed ing on many insects other than scales including aphids, small ca terpillars, whitefly, mites, thrips, psyllids, mealybugs, and other soft bodied insects and thei r eggs. These beetles are also voracious feeders of the immature stages and eggs of scale insects. The first introduction of R. lophanthae into the United States was initiated by Albert Koeble and releases occurred betw een 1889 and 1892 (Greathead 1973). It failed to control Saissetia oleae (Olivier), but did establish itself as a predator of armored scales on citrus and eventually sp read throughout the rest of the United States. It was introduced into Hawaii from California in 1894 (Heu et al. 2003) a nd has been recorded feeding on CAS (Hara et al. 2004, personal obse rvation). The larvae and adults have been observed feeding on eggs, immature, and adult CAS. This beetle was imported and officially released in Florida to control oleander scale, Aspidiotus nerii Bouch, and other

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13 scale insects after 1982 (Frank and McC oy 1994). However, many specimens in the Florida State Collection of Arthropods (FSCA) were collected in the state as early as 1936, indicating that this beetle has been established in Florida more than 70 years. Interestingly, on scaleinfested cycads in Fl orida it has been seen only in downtown Tampa and on the campus of Florida St ate University in Tallahassee. Following emergence from the egg, the small, gray larvae immediately begin to feed on scale eggs and crawlers, which are much more numerous than adult scales, but will feed on the adult scales as well. Th e larvae pass through 4 instars, growing to around 3 mm before pupation. The beetles will often pupate on the plant where the food source is found. At 25C, the total development of the life cycle of R. lophanthae takes about 34 days (Stathas et al. 2002). The adu lt beetles can live up to 9 months but average between 5 and 6 (Stathas 2000). These beetles are exceptional biological control agents because of their high fecundity, lack of parasitoids, the absence of diapause, and their resistance to low temperature especially in the immature stages (Rubs tov 1952, Smirnoff 1950, Stathas 2000). Female R. lophanthae are able to lay hundr eds of eggs in a lifetime. Stathas (2001) found that not only did he not find any parasitized beetles in the field but even when reared with known parasi toids of other coccinellids in the laboratory none of the Rhyzobius larvae or adults were parasitized. This paralleled similar findings in nature (Rubstov 1952, Smirnoff 1950). Stathas (2001) al so found that even in the winter in Greece R. lophanthae larvae could be found. In fact, both adults and larvae seem to be able to remain active at temperatures as low as 8-9C (Stathas 2000, Cividanes and Gutierrez 1996). Rhyzobius lophanthae also seems to be able to resist extreme heat,

PAGE 30

14 because Atkinson (1983) found that adult R. lophanthae had an LD 50 at about 42C. Stathas (2000) found that in Greece R. lophanthae completed 6 generations a year, while Smirnoff (1950) speculated that they may be ab le to complete as many as 7-8 generations a year in Morocco. This particular coccinellid was ment ioned by Howard (1997) as a possible biological control of CAS; but never mentioned again. Rhyzobius lophanthae has also been labeled as highly effective in c ontrolling CAS in Hawaii (Hara et al. 2004). However, there are no research data to support th is claim. In Tampa, Florida, adults and larvae have been seen in large numbers feeding on CAS (personal observation 2004). The fact that this beetle is already found in Florida makes it an appealing candidate for the biological control of CAS. Label data with specimens in the FSCA indicate this predator was captured feeding on Fiorinia theae Green, P. cockerelli Pseudaulacaspis pentagona (Targioni-Tozzetti), and Acutaspis morrisonorum Kosztarab.

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CHAPTER 2 LIFE HISTORY OF Cybocephalus nipponicus ENDRDY-YOUNGA A PREDATOR OF Aulacaspis yasumatsui TAKAGI (HOMOPTERA: DIASPIDIDAE) Introduction Cybocephalids are among the most economically important groups of natural enemies against scale insects (Alvarez and Van Driesche 1998 a). Larvae and adults are voracious predators and have many desirable traits for use as biological control agents. Cybocephalus nipponicus Endrdy-Younga was released an d later established in the Washington D.C./Maryland area to combat the euonymus scale, Unaspis euonymi (Comstock) (Drea and Carlson 1988, Drea and Hendrickson 1988). Alvarez and Van Driesche (1998a) later released and established populations of C. nipponicus in New England. This same species also was re leased, under the fals e identification of Cybocephalus binotatus (Grouvelle), into the Miami area in 1998 for control of the cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Anonymous 1998, Howard et al. 1999, Howard and Weissling 1999). While C. nipponicus and C. binotatus appear similar, they each have very distinctive male genitalia (Endrdy-Younga 1971), and C. binotatus (Fig. 2-1A) has two large black spots on the pronotum, which are absent in C. nipponicus (Fig. 2-1B). Although C. nipponicus previously was establis hed in Florida before 1998 (according to specimen label data in the Flor ida State Collection of Arthropods), its range and abundance in the state before 1998 are unknown (Smith and Cave 2006b). The cycad aulacaspis scale (CAS) is the most damaging scale found on Cycas in Florida (Hodges et al. 2003). CAS is nativ e to Thailand but is found throughout China 15

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16 and southeastern Asia, as well as on several Caribbean islands, Florida, and Hawaii (BenDov et al. 2003). In 1992, CAS had become such a problem in Hong Kong that 70-100% mortality was recorded in infested king sagos, Cycas revoluta Thunberg (Hodgson and Martin 2001) The first detection of CAS in Florida occurred in 1996 in Miami at the Montgomery Botanical Center. The scale was thought to have arrived on infested cycads imported from southeastern Asia (Howard and Weissling 1999). By the end of 1997, CAS had spread throughout Miami and as far north as Lake Okeechobee and could be found on 20 species of cycads (Howard and Weissling 1999); however it seems to prefer Cycas and Stangeria (Emshousen pers. comm. 2004). This led to the spread of CAS to Hawaii in 1998 through the legal importation of infested cycads from Florida (Hodgson and Martin 2001). At present, CAS has been reported from Pensacola east to Jacksonville and south into the Florida Keys. Infested cycads in northern Florida were suspected transplants from southern nurseries rather than natural progression of the scale northward. The objective of this study is to collect life history data on C. nipponicus using CAS as prey, and compare these to the results from Tanaka and Inoue (1980) and Alvarez and Van Driesche (1998a) using euonymus s cale as prey. A better understanding of these beetles using different prey will lead to gr eater understand ing of how they perform as biological control agents in the field, and consequently increase success in controlling CAS. Materials and Methods A colony of CAS was reared on king sago in a sealed greenhouse to keep out possible predators and/or parasitoids. Sma ll king sago specimens were infested with CAS by placing large numbers (>100) of eggs on each plant. Once infested, plants were

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17 then placed in contact with other non-infested plants to sp read non-parasitized scales to other plants. Thus, we could be confident th at parasitoids and predators did not invade the colony. A colony of C. nipponicus was initiated from individuals collected in south Miami (N25 W80). Aphanogmus albicoxalis Evans and Dessart (Hymenoptera: Ceraphronidae) parasitizes the pupae of C. nipponicus in southern Florida (Evans et al. 2005), but this parasitoid was excluded by coll ecting only adult bee tles and subsequently rearing future generations in sealed cages (0.5m x 0.5m x 0.5m). Cy cad leaves infested with CAS were cut from the colony plant, with rachis bases placed in floral water tubes for hydration, and then placed in rearing cages. The leaves and interior of the cage were misted with distilled water every three da ys. A moist sponge also was provided for hydration. Leaves were replaced every three weeks. Old leaves were held in separate cages for three weeks to recover emerging bee tles. Beetle and scale voucher specimens were placed in the Florida State Co llection of Arthropods (FSCA). All life cycle studies were carried out in temperatureand humidity-controlled cabinets set at 25 C with a relative humidity of 80% and a photoperiod of 14:10 (L:D). Each treatment was initiated by isolating 2530 mating pairs of beetles randomly selected from the laboratory colony. Each pair was placed in a 25-dram plastic vial with one C. revoluta leaflet infested with male and female CAS. After 24 hours the beetles were removed and eggs collected from beneath the scale armor on the leaflet. Eggs were measured (length and width), and then, to simulate natural conditions, placed on the surface of a small, clean piece of C. revoluta leaflet and covered with the armor of an adult female CAS. The leaflet piece was placed in the well of a rectangular tissue culture

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18 dish, covered with parafilm, and placed in the environmental chamber. Eggs were checked daily until larvae emerged. Newly emerged larvae were placed in 25-dram plastic vials with freshly-cut, CASinfested leaflets on top of sterilized sand in the bottom of the vial. Old leaflets were replaced with fresh, infested leaflets every five days throughout the life cycle. A fine mesh cloth was placed over the top of the vials to allow airflow. Larval and pupal development was checked daily. Upon emergence from the pupal case the adults remained in plastic vials and were provided with 20-30 fresh CAS every 3 days. Adults were checked daily until death. Eggs, larvae, and pupae were critical-point dried in a Tousimis samdri-780 A, and sputter coated with a gold-palladium a lloy. Images were taken with a JEOL JSM5510LV scanning electron microscope and a Syncroscopy automontage photography system. Descriptive statistics were generated in SAS (2001) using a PROC UNIVARIATE analysis. The dependent variab le was number of days in ea ch stage and the independent variable was the stage itself. PROC UNI VARIATE and PROC t-test were used to generate statistics for adult longevity. Results and Discussion The egg of C. nipponicus is elongate oval with both e nds rounded, relatively large measuring 0.42 mm by 0.20 mm (n =50) and usually light gray to purple. Eggs were usually found singly inside the vacated tubular cover of a male scale or under the armor of a female scale, usually with a live scale beneath but occasionally with a dead female. During low scale density, as many as 5 eggs under one female armor were observed. An egg deposited within the male scale cover fit snugly and had almost the same diameter as

PAGE 35

19 the scale cover, thereby allowing only one e gg to be placed in one male cover. The surface was smooth aside from debris sticking to the surface (Fig. 2-2) and was slightly tacky, allowing the egg to stick to the substrate. Females ty pically laid about 3 eggs per day and on average 288 eggs in a lifetime (Alvarez and Van Driesche 1998a). Eggs hatched about 7 days after oviposition (Table 2-1). Eyespots coul d be seen 1-2 days before larval emergence. When the larva emerged, the chorion split along the longitudinal axis and the larva wriggled free. This process took about 15 minutes compared to the 30 to 45 minutes reported by Blumberg and Swirsk i (1982) on the life history of Cybocephalus micans Reitter and Cybocephalus nigriceps nigriceps (J. Sahlberg). The neonate larvae were white or yellowish with long setae along the body, but after f eeding for a day turned light purple or lavender, with 4 black stemmata on each side of the head (Fig 2-3A, B). Not only were larvae covered in long, slender set ae but also shorter trumpet-shaped setae (Fig. 2-4). After emergence, larvae immediately began to fe ed either on the scale eggs sharing the space beneath the armor or rarely on the female scale. If an egg hatched in a male scale cover, the larva would go to th e nearest food source. Larvae continued to move from scale to scale feeding on males, females, and eggs but spent the most time underneath female armor. They also were seen cannibalizing othe r larvae when scale density was extremely low, as mentioned by Alvarez and Van Driesche (1998a). Larvae fed for 9 to 10 days. Three instars (Fig. 2-5A, B) were observed, similar to C. micans and C. n. nigriceps (Blumberg and Swirski 1982). However, Ahmad (1970) recorded four instars in Cybocephalus semiflavus Champion. When molting, the cuticle ruptured along the top

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20 of the head capsule and along the median dorsal region of the body. The larva emerged from the anterior portion of the old exuviae firs t, and then wriggled vigorously to extract the posterior body portion. The posterior portio n of the old exuviae remained attached to the substrate. Average larval developm ent was about 14 days (Table 2-1). If bright light was shone on larvae they immediately moved underneath a scale armor or debris. Once disturbed larvae raised the head and body away from the leaf surface, arching the body into a C-shape, a nd holding to the substrate with the conical protuberance found on segments 8 and 9. This thr eat posture is similar to that used by the larva to extract themselves from old exuviae. Larvae became less active 2 to 3 days prior to pupation, stopped feeding, eventually becoming immobile and attaching the posterior of the abdomen to the substrate before forming the pupal case. Larvae gathered pieces of scale armor and incorporated these pieces into an ovoid pupal chamber with one end flatter where it attached to the substrate (Fig. 2-6A). Pupal chambers often were found in the an terior portion between leaflet and leaf rachis or near the leaflet base. However, pupal chambers also were observed along leaflet and rachis. When not given access to scales, larvae used sand or other organic material to construct the pupal chamber (Fig. 2-6B), sometimes attaching to the leaf or dropping to the sand. Infrequently, larvae dr opped to the sand and made a sand cocoon even when scales were available. This behavior is common in Cybocephalus and has been suggested or recorded for other sp ecies (Clausen and Berry 1932, Flanders 1934, Smirnoff 1954, Blumberg and Swirski 1982). All pupae exhibited typical exarate

PAGE 37

21 features (Fig. 2-7A, B). Pupa tion lasted about 18 days (Tab le 2-1). Beetles exited the chamber by chewing an emergence hole. There was no significant difference in developmental time from egg to adult between sexes (t=1.50, df=52, P=0.1389). Total development from egg to adult lasted about 40 days (Table 2-1), thus it is concei vable that 7-8 generati ons could be produced per year in southern Florida or other areas with amenable temperatures. Alvarez and Van Driesche (1998a) suggested that these beetles were capable of producing 3 generations per year in New England. Preoviposition lasted about 4 days (n=29) with some females laying eggs as early as 2 days (n=2) after emergen ce. Adults (Fig. 2-8) began feeding soon after emergence and consumed an average of 4 scales per day, with females typically eating more than the males. Disproportionate feeding is probably due to size, because males are smaller than the females. Maximum female longevity was 190 days, with an average of about 110 days (Table 2-1). Average longevity of ma les was 89 days (Table 2-1) with a maximum of 155 days. Due to high variation, a t-te st showed no significant difference between longevity of the sexes (t=1.50, df=52, P= 0.1389). These findings are intermediate between the 78 days for males and 99 days for females reported by Alvarez and Van Driesche (1998a) at 22 C and 122 days for males and 143 days for females reported by Tanaka and Inoue (1980), who do not detail their experimental temperatures used. Shorter male longevity may be due to thei r more active nature. Males were observed moving around the cages more of ten than females. Often, 5 or 6 males would chase a female for hours before finding a new female to pursue. The sex ratio of emerging adults was 23:31 (male:female).

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22 Our results were similar to those of Al varez and Van Driesche (1998a) and Tanaka and Inoue (1980). However, all stages developed slower in the former study (9.1 days for eggs, 14.5 days for larvae, and 20.4 days for pupae) in comparison to our results, but this maybe due to their lower experimental temp eratures or the food source. Population differences may account for some discrepancie s in the life history parameters of the beetles in each study. The beetles studied by Tanaka and Inoue (1980) originated in Japan and those studied by Alvarez and Van Driesche (1998a) originated in Beijing, China. Our beetles may have originated in Thailand, however, this is only speculation considering the beetles were present in Florida before the recorded introduction in 1998 (Smith and Cave, in press). Mortality rate s in our study also were slightly higher, possibly due to high humidity. Fungal growth was a consistent problem in the humid environment of our rearing chambers. Scales often became so encrusted on the cycad leaflet that saprophytic fungi quickly spread. Alvarez and Van Driesche (1998a) did not record relative humidity, thus a comparison cannot be made. Mortality was highest during the larval st age. The first few days of larval development proved to be most difficult (Fig. 2-9). Eggs and pupae seemed to be quite hardy with mortality of 22% and 9%, respectiv ely (Table 2-2), which are similar to the 14% and 8% mortality rates found by Alvarez an d Van Driesche (1998a). Therefore, the 86% mortality rate in the larval stage (Tab le 2-2) is interesting and not predicted. However, when reared on San Jose scale, Quadraspidiotus perniciosus (Comstock), Alvarez and Van Driesche (1998a) found the larval stage of C. nipponicus had a correspondingly high 77% mortality rate.

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23 Certain aspects of the biology and life history of other Cybocephalus species have been studied, typically as part of biological control projects, e.g., Cybocephalus rufifrons Reitter (De Marzo 1995), Cybocephalus freyi Endrdy-Younga (Lupi 2003), and Cybocephalus fodori Endrdy-Younga (Katsoyannos 1984) ha ve been studied in Europe; and Cybocephalus semiflavus Champion (Ahmad 1970) and Cybocephalus gibbulus Erichson (Nohara and Iwata 1988) have been studi ed in Asia. In the Middle East, several studies have been carried out on C. aegyptiacus, C. binotatus C. micans and C. n. nigriceps (Blumberg 1973, 1976; Blumberg and Sw irski 1974 a,b, 1982). In Australia, Kirejtshuk et al. (1997) described Cybocephalus aleyrodephagus and studied its life cycle. The life histories of these species do not differ dramatically from C. nipponicus and there seems to be fairly consiste nt life cycle and feeding habits. In the absence of prey, female cybocephalid s are able to withhold eggs for up to 2 days, indicating that oviposition strategy is not only governed by food consumption but also by qualitative features of the scale popul ation (Alvarez and Van Driesche 1998b). In the presence of high scale density, cybo cephalids increase egg production until an asymptote is reached. Again, this allows c ybocephalids to maintain their populations at low scale densities. If circumstances allo w, female cybocephalids will lay only one egg under a single scale cover. However, if the number of scales in a patch is low, and the beetle cannot find new scales, it will lay eggs under a scale under which eggs are already present (Alvarez and Van Driesc he 1998b). The beetle larvae are forced to consume more prey when feeding on younger scales, ther efore for greater offspring survivability female cybocephalids prefer oviposition on olde r scales that provide a larger food source and require less searching (Alvarez and Van Driesche 1998b). Alvarez and Van Driesche

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24 (1998b) found that the highest larval survival rate could be found in larvae feeding on scales older than 30 days. Blumbe rg and Swirski (1982) noted that C. n. nigriceps rarely laid eggs under dead female scales. Evans et al. (2005) described two species of Aphanogmus that parasitize C. nipponicus pupae. Aphanogmus inamicus Evans and Dessart emerge d in quarantine from hosts collected in Thailand, and A. albicoxalis occurs naturally in Fl orida. The latter species is broadly distributed in southern Fl orida, with collection sites in Collier, MiamiDade, and St. Lucie counties. The levels of parasitism in C. nipponicus populations in Florida are currently undetermined, but large numbers of these wasps occasionally have been seen on cycads. Cybocephalus nipponicus will probably always be considered a supplementary predator for the control of CAS. However, some attributes make these beetles very attractive as biological control agents, including a long lifespa n, some ability to resist or protect themselves from pesticides (Alv arez and Van Driesche 1998a, Katsoyannos 1984, Kehat et al. 1974), and most importantly, the ability to persist at low scale densities (Alvarez and Van Driesche 1998b). However, even in combination with the parasitoid Coccobius fulvus Compere and Annecke (Hodges et al. 2003) these beetles are unable to adequately control populati ons of CAS in Florida.

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25 Figure 2-1. Dorsal habitus. A, Cybocephalus binotatus B, Cybocephalus nipponicus Figure 2-2. Egg of Cybocephalus nipponicus

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26 Figure 2-3. First instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral habitus. Figure 2-4. Short trumpet-shaped setae on third instar larva of Cybocephalus nipponicus.

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27 Figure 2-5. Third instar larva of Cybocephalus nipponicus A, Dorsal habitus. B, Ventral oblique habitus.

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28 Figure 2-6. Pupal chambers of Cybocephalus nipponicus. A, Pupal chamber made from female scale covers. B, Pupal chamber made from sand.

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29 Figure 2-7. Pupa of Cybocephalus nipponicus A, Dorsal habitus. B, Ventral habitus.

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30 Figure 2-8. Adult C. nipponicus, lateral habitus. 0 5 10 15 20 25 30 123456789101112131415 Age in Days% Mortality Figure 2-9. Larval mortality over time.

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31 Table 2-1. Development of Cybocephalus nipponicus on Aulacaspis yasumatsui. Stage Mean (Days) SE Range (min-max) (Days) N Egg 7.3 0.1 4 (5-9) 131 Larva 13.7 0.1 4 (12-16) 94 Pupa 18.6 0.2 6(16-22) 44 Total development time 39.5 Male life-span 89.1 9.7 146 (9-155) 23 Female life-span 110.0 9.6 175 (15-190) 31 Table 2-2. Mortality (%) of immature stages of Cybocephalus nipponicus during rearing on Aulacaspis yasumatsui Stage (x) No. Alive at Start (l x ) No. Dying in Stage (d x ) Apparent Mortality (q x ) Egg 200 44 0.22 Larva 156 134 0.86 Pupa 22 2 0.09 Adult 20

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CHAPTER 3 PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus AND Rhyzobius lophanthae (COLEOPTERA: CYBOCEPH ALIDAE, COCCINELLIDAE) Introduction Beetles of the families Coccinellidae and Cybocephalidae are the most economically important groups of predators of diaspidid scales in the world (Blumberg and Swirski 1982). Cybocephalus nipponicus Endrdy-Younga (Cybocephalidae) and Rhyzobius lophanthae Blaisdell (Coccinellidae) are commo nly used as biological control agents for many armored scale pests. Rhyzobius lophanthae has been established in Florida since the 1930s (according to specimen label data in the Florida State Collection of Arthropods). Cybocephalus nipponicus, misidentified as Cybocephalus binotatus Grouvelle, was recently released in south Florida in an effort to control the cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Anon. 1998; Howard et al. 1999; Howard and Weissling 1999). CAS is the mo st economically damaging scale to cycads that the state of Florida has ever seen (Hodges et al. 2003). Although C. nipponicus is present in Hawaii (Heu and Chun 2000), R. lophanthae is usually sugges ted as the better control agent of CAS (Heu et al. 2003; A. Hara, pers onal communication). In both places, CAS has continued to spread and multiply. A more promising approach to controlling CAS would be one us ing integrated pest management (IPM). In this manner, a combination of pesticides and biological control would be used to combat CAS. There has been some success controlling CAS with various pesticides. Oils, either an ultra-fine horticultural oil or a product containing fish oils, seem to be the most 32

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33 effective chemical control method (Hodges et al 2003). This is not surprising given that oils have long been used to control armored scale insects. The oil not only covers the insects and suffocates them but also covers the surface of the plant making it difficult for crawlers to settle onto the plant (Howard and Weissling 199 9). Soaps are quite popular with homeowners; but they must be applie d frequently, in some cases once a week (personal observation). The effective application of pestic ides for control of CAS is difficult due to the scales tendency to heavily infest the abaxial surface of leaves, which is difficult to spray (Howard and We issling 1999). In the case of C. revoluta the architecture of the plant itself, with the marg ins of the leaflets curling down and inward to form an arch on the abaxial surface of the l eaflet, makes foliar treatments inefficient (Hodges et al. 2003). Frequent or as needed ap plications of oils seems to be the most effective technique for controlling CAS, and by mixing oil with contact pesticides such as malathion, even greater scale mortality can be achieved (Hodges et al 2003). The use of systemic pesticides such as dimethoate a nd contact pesticides like methidathion has yielded mixed results, being very effective in some instances and completely ineffective in other cases (Hodges et al. 2003). Imidacloprid used as a soil drench can be very effective, but Howard and Weissling (1999) found that this product had to be mixed at very high concentrations to be effective. Th is product can also be used as a foliar spray. The reproductive biology of C. nipponicus makes it a good biological control agent. Alvarez and Van Driesche (1998a ) found that, at low scale densities, C. nipponicus was able to maintain its populations and maintain populations of euonymus scale, Unaspis euonymi (Comstock), and San Jose scale, Quadraspidiotus perniciosus (Comstock), in check. In the presence of greater scale densities, C. nipponicus will

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34 increase its egg production accordingly. With a total life cycle from egg to adult only taking around 44 days (Smith and Cave 2006b), it is conceivable th at 5-6 generations could be produced every year in Florida. Cybocephalus nipponicus is available commercially in the U.S. market. Rhyzobius lophanthae is an exceptional biological control agent because of its high fecundity, lack of parasitoids, the absence of diapause, and resistance to low temperatures especially in the immature stages (Rubstov 1952; Smirnoff 1950; Stathas 2000). Female R. lophanthae are able to lay hundreds of eggs in a lifetime (Stathas 2000). Rhyzobius lophanthae also seems to be able to resist extr eme heat. Atkinson (1983) found that adult R. lophanthae could not survive for long at 42 C. Rhyzobius lophanthae is also available commercially in the U.S. market. This study was conducted to determine the susceptibility of C. nipponicus and R. lophanthae to six pesticides commonly used in the control of CAS. Given the established presence of both predators on cycads in south Florida and their commercial availability, it is very important to learn what effects th e commonly used pesticides against CAS will have on them. This information is vital for development of IPM programs aimed at controlling CAS. Materials and Methods Insects Adult R. lophanthae were reared at and purchased from Rincon-Vitova Insectaries (Ventura, California). Adult C. nipponicus were also purchased from Rincon-Vitova but were reared by Philip Alampi Beneficial In sect Laboratory, New Jersey Department of Agriculture. Both beetle sp ecies were maintained in Plex iglas cages at 25 C prior to testing. All life stag es of CAS were provided as a food source.

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35 Food was not provided during testing because of the very small size of the beetles (1 mm and 2.5 mm in length). The beetles could have conceivably perched on the food source for long periods of time, never coming in to contact with the walls of the treated vial. Preliminary studies indicated that a 24-hour period without food would not unduly stress the beetles. On average, untreated C. nipponicus survived for 8-9 days (n=30) and untreated R. lophanthae lived for 5-6 days (n=30) before dying of starvation. Cotton used to stopper the vials was soaked in water to prevent dehydration. Bioassays Using Coated Glass Vial Method A coated glass vial method (Plapp 1971; Amalin et al. 2000; Snodgrass 1996; Snodgrass et al. 2005) was used to determin e the chemical susceptibility of adult R. lophanthae and C. nipponicus to six pesticides used to control CAS (Howard et al. 1997; Howard and Weissling 1999; Weissling et al. 1999; Hodges et al. 2003; Emshousen and Mannion 2004). This is a very effective method for testing the chemical susceptibility of small arthropods (Amalin et al. 2000) such as R. lophanthae and especially C. nipponicus because of its extremely small size. The six pesticides tested were fish oil emulsion (Organocide), insecticidal soap (Garde n Safe, Inc.), imidacloprid (Provado), malathion (Spectracide, Inc.), methidathi on (Supracide), and dimethoate (Cygon). The fish oil and insecticidal soap were purchased at commercial grade, while the imidacloprid (99% purity), ma lathion (98% purity), methid athion (98.6% purity), and dimethoate (98.7% purity) were purchased in the technical grade from Chem Service (West Chester, PA). All pesticides were dissolved in acetone, except the insecticidal soap which does not dissolve in acetone. Instea d, the insecticidal soap was di ssolved in 95% ethanol. The fish oil was shaken in a paint shaker after being placed in ace tone in order to break up the

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36 oil into fine globules. Each pesticide was se parated into three dilu tions: field rate, twice field rate, and one-half field rate. The fi eld rate was taken from label data for each pesticide as directed for use against scal e insects. A small amount (0.5 ml) of the pesticide working solution was dispensed into 20-ml scintillation vials. Concentrations of active ingredient for the working solution and the amount of active ingredient residue within the vials can be seen in Table 3-1. Vials were hand turned until the acetone or ethanol completely evaporated leaving an inse cticidal residue on the inner surface. Vials treated with only acetone or etha nol, as well as untreated vials, were used as controls. A single beetle was placed into a treated vial. All beetles ha d emerged from pupae within the previous 14 days. Vials were sealed with cotton soaked in water allowing the beetles to drink. Vials were placed upright in a ve ntilated cabinet with a fume hood and at a constant temperature of 25 C and 80% relati ve humidity for 24 hours. For each treatment of 10 beetles, 5 females and 5 males were used. Each treatment of 10 beetles was replicated 3 times for each dosage. All trials were carri ed out the same day as the pesticide was applied to the vials. Mortality of beetles was determined immediat ely after the 24-hour period. A beetle was considered dead if it was not moving or could not right itself. Percent survivorship was measured as the proportion of 30 beet les alive after a 24-hour exposure to the pesticides. Statistical Analysis All descriptive statistics were genera ted in EXCEL (Microsoft 2000). The mortality rates for each pesticide were comp ared using the Student-Newman-Keuls mean separation test (SAS Institute 2001).

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37 Results Of the six pesticides tested on adult C. nipponicus and R. lophanthae, three (methidathion, dimethoate, and malathion) cau sed >90% mortality at all concentrations, while the other three (fish oil, insecticidal soap, and imidacloprid) were less toxic but still caused very high mortality (Tables 3-2 & 3-3). Effects of Pesticide on C. nipponicus Cybocephalus nipponicus was extremely susceptible to all pesticides. The three least toxic pesticides were imidacloprid, insectic idal soap, and fish oil (Table 3-2). There were significant differences in mortality between concentrations amongst these three pesticides (Table 3-4). Fish oil was still qui te toxic to this beetle and due to its very small size, C. nipponicus would often get trapped in sma ll globules of oil, eventually dying from suffocation. Effects of Pesticide on R. lophanthae Rhyzobius lophanthae was more tolerant than C. nipponicus to the experimental pesticides, although mortality rates were also high for this species. The three least toxic pesticides to R. lophanthae were imidacloprid, insecticidal so ap, and fish oil (Table 3-3). There were significant differences in survivor ship between concentrations of these three pesticides (Table 3-4). Rhyzobius lophanthae about twice the size of C. nipponicus had much less difficulty traversing oil globul es on the surface of the vials. Discussion There was a significant difference between mort ality in the control and that of even the lowest pesticide concentration. This sens itivity to pesticides makes an IPM approach to the control of CAS quite difficult. Unfortunately, most of the success in chemically

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38 controlling CAS has involved very toxic pesticides often being used at higher than recommended doses (Howard and Weiss ling 1999, Weissling et al. 1999). The high mortalities experienced by C. nipponicus and R. lophanthae are not unexpected. Nakao et al. (1985) found that all 18 species of Co ccinellidae inhabiting Japanese citrus groves were se verely affected by the applica tion of pesticides, including methidathion and dimethoate. They also found that Cybocephalus gibbulus Erichson, one of the most common scale predators found in Japanese citrus groves, was virtually eliminated by long-term pesticide use. O ils have proven to be the most effective pesticides used against many plant-sucking pests, while maintaining the natural enemy populations. Erkili and Uygun (1997) found that oils were much less toxic to Cybocephalus fodori minor (Endrdy-Younga) and Chilocorus bipustulatus (Linnaeus) than was methidathion. In fact, they went as far as to say that methidathion should not be used in IPM programs. In natural conditions, the predatory beetle s may not likely be in contact with the pesticide for as long as the exposures in this experiment. However, C. nipponicus and R. lophanthae are uniquely suited for life in chemica lly-treated environments. Both beetle species place their eggs undernea th the scale cover and at least part of larval development takes place beneath the armored scale, allowi ng the beetles some protection from both the elements and pesticides (Smi rnoff 1950; Alvarez and Van Dr iesche 1998; Stathas 2001). In Greece, Katsoyannos (1984) found that C. fodori was able to survive in pesticidetreated fruit orchards. In da te palm plantations in Israel Kehat et al. (1974) found that, while all coccinellids in a chemically -treated plantation died, species of Cybocephalus survived.

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39 It is apparent from these tests that, for some pesticides, the lower the concentration of the pesticide the higher th e survivorship. However, th ese tests were conducted in a laboratory environment in which the test subj ects were in constant contact with the pesticide for 24 hours. A whole host of f actors, such as humidity, UV degradation, evaporation, and precipitation, will influence pesticid e activity in the field. Nevertheless, whenever possible, insecticidal soaps a nd fish oils should be used. While many homeowners use various types of soaps to treat CAS, this method requires treatment every 10 to 14 days, thus increasing exposure of the beetles to the pes ticide. If more toxic pesticides must be used, then applying them to hot spots rather than broadcast spraying may protect the scale predators from comple te annihilation. This type of selective spraying may also protect other entomophagous insect populations from being decimated (Kuznetsov 1997). The results of these laboratory experiment s yield some baseline data from which more research in the field can be conducted.

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40 Table 3-1. Field rates (1X) for each pesticide used. Insecticide Work ing Solution ( g*AI/ml) Insecticide residue ( g*AI/cm 2 ) Organocide 47000 8.29 Insecticidal Soap 512300 27.71 Imidacloprid 106 2.40 Methidathion 233 5.26 Dimethoate 305 6.91 Malathion 1990 45.07 AI = Active Ingredient Table 3-2. Percent mortality of Cybocephalus nipponicus per 30 individuals exposed. X = field rate. Pesticide % beetle mortality % beetle mortality % beetle mortality % beetle mortality at 0X at 0.5X at 1X at 2X Organocide 83 100 96 Insecticidal Soap 66 86 96 Imidacloprid 76 93 100 Methidathion 100 100 100 Dimethoate 100 96 100 Malathion 93 100 100 Control (Acetone) 0 Control (Ethanol) 0 Control (No coating) 0 Table 3-3. Percent mortality of Rhyzobius lophanthae per 30 individuals exposed. X = field rate. Pesticide % beetle mortality % beetle mortality % beetle mortality % beetle mortality at 0X at 0.5X at 1X at 2X Organocide 46 83 100 Insecticidal Soap 43 76 96 Imidacloprid 63 80 100 Methidathion 100 100 100 Dimethoate 100 96 100 Malathion 93 90 96 Control (Acetone) 0 Control (Ethanol) 6 Control (No coating) 0

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41 Table 3-4. Student-Newman-Keuls test showi ng ranked values of mortality of adult Cybocephalus nipponicus and Rhyzobius lophanthae C. nipponicus Dose Imidacloprid Organocide Insecticidal soap 0.0X 2.0A 2.0A 2.0A 0.5X 5.5B 5.3B 5.3B 1.0X 8.5C 8.6C 8.3C 2.0X 10.0C 10.0C 10.3C R. lophanthae Dose Imidacloprid Organocide Insecticidal soap 0.0X 2.0A 2.0A 2.0A 0.5X 5.6B 5.0B 5.3B 1.0X 7.3B 8.0C 7.8C 2.0X 11.0C 11.0D 10.8D Mean ranks within columns with the same letter are not significantly different (P=0.05).

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CHAPTER 4 THE CYBOCEPHALIDAE (COLEOPTERA) OF AMERICA NORTH OF MEXICO Introduction The family Cybocephalidae consists of seven genera: Cybocephalus (Erichson 1844), Endrodiellus (Endrdy-Younga 1962a), Hierronius (Endrdy-Younga 1968), Horadion (Endrdy-Younga 1976), Pastillodes (Endrdy-Younga 1968), Pastillus (Endrdy-Younga 1962a), and Pycnocephalus (Sharp 1891). By far the largest of these is Cybocephalus, which contains more than 150 described species found throughout the world (Tian 2000, Yu and Tian 1995). The Cybocephalidae differ in many ways from the Nitidulidae. They are predatory, almost exclusively feeding on scale insects, while nitidulids are known for feeding on decaying plant material and fruits, plant sap, fungi, and occasionally pollen and honey. The morphology of the Cybocephalidae is al so quite different from that of the Nitidulidae. Cybocephalid adults have a 4-44 tarsal formula instead of 5-5-5 found in Nitidulidae. There are 5 visibl e ventral plates (leaving out the male anal plate) and 5 abdominal spiracles in cybocephalids instead of the 6 and 6 that occur in nititdulids. The body of cybocephalids is retractile allowing th e mandibles in repose to rest against the metasternum, unlike any other nitidulid. The larvae of Cybocephalidae have a head without dorsal sutures, lack pregomphi and urogomphi on abdominal tergite XI, and have hypostomal rods with divergent hypostoma l ridges present posteriorly, hypopharynx without a sclerome and bracons, maxillae wi thout mola, and annular spiracles with 2 lateral air tubes. In contrast, the larvae of Nitidulidae have pregomphi and urogomphi, no 42

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43 hypostomal rods but with hypostomal ri dges strongly convergent posteriorly, hypopharynx with a sclerome and bracons, max illae with raised mola, and biforous spiracles (Kirejtchuk 1997). These larval diffe rences were also illustrated by Bving and Craighead (1931) and Hayashi (1978). In sp ite of this, the group has been shuffled between family (Jacquelin du Val 1858, Murray 1864, Bving and Craighead 1931, Parsons 1943, Smirnoff 1954, Endrdy-Younga 1968) and subfamily (Erichson 1844, Horn 1879, Grouvelle 1913, Kirejtshuk 1997, Habeck 2002) status. We chose to recognize the Cybocephalidae as a family in accordance with the work of the world expert in this group, Se bastian Endrdy-Younga. The adults of Cybocephalus are often confused with those of Clambidae, Phalacridae, and sometimes Coccinellidae ( Microweisea Cockerell and Gnathoweisea Gordon). The major feature that distinguishes members of Cybocephalus from all of these families is their extremely broad head. The head of Cybocephalus is almost as wide as the pronotum, unlike that of all the aforementioned families. Adult clambids have extremely enlarged hind coxae, a clypeus th at entirely covers the mouthparts, and a 2segmented antennal club, whereas cybocephali ds have small hind coxae, a relatively short clypeus and a 3-segmented antennal club. Phalacrids have a much more elongate club than cybocephalids and a 55-5 tarsal formula rather than the 4-4-4 formula found in cybocephalids. Gnathoweisea has an extremely long head compared to a short, wide head in cybocephalids. Also, both Gnathoweisea and Microweisea have the head inserted into the prothorax, unlike cybocephali ds which have the head completely outside the pronotum, and both genera have distinct ive tarsomeres easily distinguishing them from the Cybocephalidae.

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44 Cybocephalids are primarily known for feed ing on armored scales (Diaspididae) (Flanders 1934, Clausen 1940, Vinson 1959, Endrdy-Younga 1968, Blumberg and Swirski 1974a,b, Rosen and DeBach 1978, Alvar ez and Van Dreische 1998a). However, they have also been reported feeding on whiteflies (Aleyrodidae) (Clausen and Berry 1932, Chandra and Avasthy 1978, Swirski, et al. 1987, Kapadia and Puri 1993, Kajita et al. 1991, Kirejtshuk et al. 1997, Ramani 2000, Tian and Ramani 2003), mealybugs (Pseudococcidae) (Endrdy-Younga 1982), and citrus red mite, Panonychus citri (McGregor) (Tanaka and Inoue 1980). These minute beetles can be found throughout the world, but relatively few of them have been studied. Grouvelle (1913) described 5 species of Cybocephalus from the Seychelles, while Vinson (1959) found at le ast 8 new species in addition to the two known species of Cybocephalus from the small Mascarene Islands. More recently, Tian and Peng (1997) recorded at least 8 species of Cybocephalus from Hainan Island in China. It seems very unlikely that all of th ese island groups, with such a finite amount of land area, would contain such rich Cybocephalus diversity while larger land masses would not. Considering that th ere are 150 described species of Cybocephalus, these studies would seem to indicate that these bee tles are often overlooked or simply ignored. Cybocephalus species are particularly difficult to identify because they are very small (0.5-2.5 mm.), compact, extremely convex and have a deflexed head and the ability to retract their appendages. For this reason, descriptions of male genitalia are an important accompaniment to any description or key to species within this genus (Vinson 1959). Male genitalia in conjunction with othe r useful diagnostic characters such as the form of the antenna can make identification less difficult.

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45 The objectives of this study are to 1) develop techniques for the identification of the cybocephalids of America north of Mexico; 2) validate and revi ew the identification of adventive cybocephalids released in Florida; 3) describe new North American species; 4) determine major prey species of th ese cybocephalids; a nd 5) report the known distribution of each species in America north of Mexico. Because some species of Cybocephalus are being released in several countries as biological control agents, it is absolutely imperative that a clear and concise form of identification be devised. Taxonomic suppor t is especially impor tant because correct identification of pests and their natural enemies are [sic] absolutely essential for effective biological control (Van Driesche and Bellows 1993). Taxonomic History The genus Cybocephalus was described by Erichson in 1844, who also described five species from Europe, Africa, and Asia LeConte (1863) described the first New World cybocephalid, Cybocephalus nigritulus Horn described a second species, Cybocephalus californicus from North America in 1879. Since that time no new species of Cybocephalus have been described from America north of Mexico. The only other species known to occur in this region is Cybocephalus nipponicus Endrdy-Younga, which was brought from Asia as a biological control agent in the late 1980s (Drea and Carlson 1988). Most taxonomic research on this group was carried out by EndrdyYounga (1962a,b, 1963, 1964a,b, 1965, 1967a,b, 1968, 1969, 1971a,b, 1976, 1979, 1982, 1984) on beetles found in Africa, Eurasia, and Oceania. Lately, an enormous amount of work has been done on the cybocephalids of China and southern Asia by M. Tian and coworkers (Tian 1995, 1996, 2000; Tian and Pang 1994; Tian and Ramani 2003; Tian and Yu 1994; Tian and Peng 1997; Yu, 1994, 1995a,b; Yu and Tian 1995).

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46 Materials and Methods Materials For this study, 871 specimens belonging to the genus Cybocephalus were examined. The holotype of C. nigritulus the lectotype of C. californicus and a topotype of Cybocephalus binotatus (Grouvelle) were included. Specimens were borrowed from the following institutions and private collectio ns; names of the curato rs and owners are in parentheses: AAIC Albert Allen Insect Coll ection, Boise, ID, (Albert Allen) BMNH Natural History Museum, London [formerly British Museum (Natural History)], UK, (Maxwell Barclay) CSCA California State Collection of Arth ropods, Sacramento, CA, (Chuck Bellamy) CNCI Canadian National Collection of In sects, Ottawa, Ontario, (Anthony Davies) EMEC Essig Museum of Entomology, University of California, Berkeley, CA, (Cheryl Barr) FSCA Florida State Collecti on of Arthropods, Gainesville, FL, (Paul Skelley and Michael Thomas) HGIC Holly Glenn Insect Collection, Homestead, FL, (Holly Glenn) INHS Illinois Natural History Survey, University of Illinois, Champaign, IL, (Colin Favret) LACM Los Angeles County Museum of Natural History, University of California, Los Angeles, CA, (Weiping Xie) LSAM Louisiana State Arthropod Museum, Louisiana State University, Baton Rouge, LA, (Andrew Cline a nd Chris Carlton) MTEC Montana Entomology Collection, Montana State University, Bozeman, MT, (Michael Ivie) MCZC Museum of Comparative Zoology Collec tion, Harvard University, Cambridge, MA, (Philip Perkins)

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47 OSUC Ohio State University Collection, Muse um of Biological Diversity, Columbus, OH, (Norman Johnson) OSAC Oregon State Arthropod Collection, Oregon State Univ ersity, Corvallis, OR, (Jason Leathers) SBMN Santa Barbara Museum of Natural History, Santa Barbar a, CA, (Michael Caterino) SEMC Snow Entomological Museum, University of Kansas, Lawrence, KA, (Zack H. Falin) TAMU Texas A&M University, Department of Entomology, College Station, TX, (Ed G. Riley) TRSC Trevor Randall Smith Collection, Gainesville, FL, (Trevor Randall Smith) UCDC University of California Davis, R. M. Bohart Museum of Entomology, Davis, CA, (Steve L. Heydon) UCRC University of Californi a Riverside, Entomology Research Museum, Riverside, CA, (Doug Yanega) UCFC University of Central Florida Insect Collection, University of Central Florida, Orlando, FL, (Stuart Fullerton) WFBM W. F. Barr Entomological Museum, University of Idaho, Moscow, ID, (Frank Merickel) USNM United States National Museum, Smith sonian Institute, Washington D.C., (Gary Hevel) Methods If possible, genitalia were removed using minuten pins glued to wooden applicator sticks. The minutens were bent and twisted into whatever shape tools were necessary. While it is possible to remove only the genital plate and then extract male genitalia, this technique is extremely difficult and time -consuming. All dissections took place in glycerine due to the convex body form of Cybocephalus beetles. Typically, the abdomen was separated from the rest of the body and the entire aedeagus (Fig. 4-1) removed from the abdomen. This technique leaves most of the specimen intact, and the removed

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48 abdomen can be glued to the point behind th e specimen. The tegmen and median lobe are compressed and held together by the tegminal (lateral) struts. To separate these two parts, a minuten was wedged between the median lobe and the tegminal strut breaking one lobe of the tegminal strut, and separati ng the two pieces. In some cases, the internal sac can be removed, and the tegmen can be moved backwards sliding both tegminal struts over the posterior portion of the median lobe as well as the median strut, effectively separating the two parts without damaging the tegminal struts, the median strut, or the dorsal piece of the tegmen. The median lobe a nd tegmen (basal plate) can then be slidemounted or, to avoid distortion, mounted on a point in dimethyl hydantoin formaldehyde (DMHF). This solution is wate r soluble and dries clear. If possible, genitalia should be mounted on very shallow depression slides to avoid distortion (Fig. 6, 19, 32). This is especially true of a median lobe w ith a large and raised median plate. Specimens were cleared in 10% KOH at 24 C in preparation for disarticulation. After 24 hours, beetles were sufficiently cleared and softened for dissection. Due to their convex body form and very small size, specimens were disarticulated in glycerine. All disarticulated parts, including genitalia, were then washed in 95% ethanol and mounted on microscope slides using H oyers solution. Label data were copied onto slides verbatim with label breaks indicated by a slash ( / ). Definitions Median lobe (Fig. 4-1): Also referre d to as penis (Endrdy-Younga 1968, 1971a, 1971b; Kirejtshuk et al. 1997; Lupi 2003; Yu 1995a, 1995b). Basal plate (Fig. 4-1): This is a referen ce to the basal plate of the tegmen (EndrdyYounga 1968, 1971a, 1971b; Lupi 2003; Yu 1995a, 1995b).

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49 Cybocephalus Erichson 1844 Cybocephalus Erichson 1844: 441-442. Redescription. Form: Ovate and very convex; body contractile (Fig. 4-2). Head: Broad and deflexed. Labrum emarginate. Epistoma slightly prolonged at middle. Mandibles in repose resting against metast ernum, acute at tip with a small tooth posteriorly. Maxillae with one lobe. Antennae slightly l onger than width of head, antennal club flat with 3 antennomeres, ante nnal grooves small and convergent. Scape large and round. Thorax: Pronotum margined at base, coveri ng base of elytra, sides very short. Prosternum acutely carinate in fr ont, not prolonged behind the procoxae, procoxal cavities open behind. Mesosternum broad, oblique. Metasternum protuberant and clothed in hairs. Both the mesoand meta sternum are impressed for the reception of the middle and hind legs. Scutellum: Large, triangular. Elytra: Covering or nearly covering tip of the abdomen, apices curved. Abdomen: Five visible ventral plates (omitting the small male anal plate), and 5 abdominal spiracles. Legs: Tibiae simple, tarsi fourjointed, each tarsomere slightly dilated vent rally, second and third tarsomeres bilobed, claws simple. Median lobe: Trunk heavily sclerotized and dorsoventrally compressed. Tegmen: No parameres. Key to the species of Cybocephalus of America north of Mexico 1. Antennal club without a serrated margin (Fig. 4-12); scutellum with slightly concave margins (Fig. 4-11)................................................. kathrynae new species 1. Antennal club with a serrated margin (Fig. 4-4); scutellum with straight or slightly convex margins (Fig. 4-10)......................................................................................2 2(1) Terminal antennomere of the antennal club rounded (Fig. 4-30).............................. .............................................................................. randalli new species 2 Terminal antennomere of the antennal club truncate (Fig. 4-17) or slightly emarginate (Fig. 4-3, 4-4)........................................................................................3

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50 3(2). Antennomere 3 slightly shorter than 4 and 5 combined; male bicolored with head, prothorax, and mesosternum yellow or ta n, rest of body black (Fig. 4-29); basal plate coming to a rounded point (Fig. 4-26); median lobe as in Figs. 4-24, 4-25..... ..................................................................................... nipponicus Endrdy-Younga 3. Antennomere 3 as long or longer than 4 and 5 combined; male with head, prothorax and mesothorax black or dark brown......................................................4 4(3). All legs and antennae brown or black; elytral apic es with a large and wide impunctate area of yellowish translucen ce; male basal plate evenly rounded, without a protuberance (Fig. 4-8); me dian lobe as in Figs. 4-5-4-7.......................... ....................................................................................................... californicus Horn 4. At least profemora yellow or amber and usually antennae amber or tan; translucent impunctate area at the apices of elytra much smaller; male basal plate with a protuberance in the middle (Fig. 4-21) median lobe as in Figs. 4-18-420...... ..................................................................................................... nigritulus LeConte Cybocephalus californicus Horn (Figs. 4-3-4-9) Cybocephalus californicus Horn 1879: 320-321. Diagnosis. Male and female are black, brown, or aeneous. Antennal club is smaller than the eye and truncate or slightly emarginate at the apex, unlike the 11 th antennomere of C. randalli, which is rounded. Each antennomere of the club is distinctly separated, forming a serrated margin, unlike C. kathrynae which has a smooth club margin. Apices of the elytra have a large, wide area of yellowish transluc ence that is without punctation, which distinguishes this species from C. nigritulus. In males, the basal plate and median lobe are easily distinguished from those of all other species. Redescription. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 0.95-1.30 mm (measured from apex of clypeus to apex of elytra); breadth: 0.85-1.20 mm (measured at base of elytra). Color: Head, thorax, elytra and underside black with surface sometimes aeneous, lateral margin of pronotum and

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51 posterior margin of elytra yellowish and tr anslucent; legs and antennae brown or black. Head: Broad and convex, clypeus moderately produ ced, narrow, and slightly reflexed. Eyes large, oblong, with internal margins distinct. Ge nae not visible from above but slightly explanate when viewed late rally. Dorsal surface smooth under high magnification, distinctly alutaceous and finely punctate. Antennae 11-segmented including a club with 3 ante nnomeres, club length about size of the eye. Club flat and distinctly separated from funicle and w ith a distinctly serrated margin. First club antennomere wider than long, second club antenn omere larger than either first or third club antennomere and about as long as wide Terminal club antennomere truncate (Fig. 4-3) or emarginate (Fig. 4-4) setose and about as long as wide. Antennomere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior. Surface distinctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides slightly sinuous and ap ices rounded. Length slightly shorter than combined width (30:38). Uniformly punctate along dorsal surface, smooth at sides and base with a large impunctate area at apices of elytra. Appearing alutaceous under high magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering the surface. Legs: Femora glossy, broad, flattened and sparsely covered with short hairs. Proand mesofemora about the same width throughout le ngth. All tibiae slightly but distinctly curved and dilated towards apex. The protibiae with short hairs along outer margin.

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52 Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides parallel or slightly divergent curving into a point with declivous sides from apex (Figs. 45, 4-6). In profile, strongly curved from middle (Fig. 4-7). Median plate on surface elevated. Basal plate: Sides parallel at base, evenly rounded at apex (Fig. 4-8). Female. Nearly identical to male. Geographic distribution. Arizona, British Columbia, California, Colorado, Idaho, Montana, Nevada, New Mexico, Oklahoma, Or egon, Texas, Utah, Washington (Fig. 4-9). Hosts. This species is known to feed on or was found in association with Aonidiella aurantii (Maskell) (from label data), Chionaspis pinifolia (Fitch) (from label data), Diaspidiotus perniciosus (Comstock) (Heintz 2001), Diaspis echinocacti (Bouch) (from label data), Diaspis manzanitae Whitney (from label data), Ehrhornia cypressi (Ehrhorn) (Flanders 1934; Kartman 1946; Clausen 1940), Lecanium corni Bouch (Heintz 2001), Lepidosaphes beckii (Newman) (from label data), Mercetaspis halli (Green) (Kartman 1946), and Parlatoria blanchardi Targioni Tozzetti (from label data), all of which belong to the family Diaspididae. It has al so been seen in association with Phoenicococcus marlatti Cockerell (from label data) belonging to the family Phoenicococcidae, and some species of aleyrodids (from label data). Type material examined The so-called lectotype in the MCZC is a male specimen glued to a point with the following labels: Cal (printed) [small white rectangular label with the edge of one side dipped in purple ink] / LectoTYPE (printed) 3216 (handwritten) [red rectangular label] / Horn Coll H (printed) 3751 (handwritten) [white rectangular label] / C. californicus Cr. (handwritten) [white rectangular label] /

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53 MCZ Type (printed) 7970 (handwritte n) [red rectangular label] / Cybocephalus californicus Horn Det: Trevor Smith (printed) [wh ite rectangular label]. While this specimen is a syntype, it is not a valid lectotype because this designation was never published. This is consistent with many of Horns type specimens that had lectotype labels placed on them around 1930 without co rresponding publications. This occurred when the Horn collection was located at th e Academy of Natural Sciences, Philadelphia (Philip D. Perkins 2005, personal communicatio n). Two female specimens, labeled as paratypes in the MCZC, were also examin ed. The first paratype has the following labels: Cala (printed) (handwritten) [small white rectangul ar label] / Para-Type (printed) 3216.4 (handwritten) [blue rectangular label] / Horn Coll H (printed) 3751 (handwritten) [white rectangular label] / Type 7970 (handwritten) [red rectangular label] / Cybocephalus californicus Horn Det: Trevor Smith (printed ) [white rectangular label]. The second paratype label has the following la bels: Cal (printed) j (handwritten) [small white rectangular label] / Para-Type (pri nted) 3216.3 (handwritten) [blue rectangular label] / Horn Coll H (printed) 3751 (handwritt en) [white rectangular label] / Type 7970 (handwritten) [red re ctangular label] / Cybocephalus californicus Horn Det: Trevor Smith (printed) [white rectangular label]. Tw o male syntypes in the MCZC were also examined. The syntypes are both glued to a si ngle card with the following labels: Type (printed) 7970 (handwritten) [orange rectangular label] / Cybocephalus californicus Horn (handwritten with Cr. crossed and Horn written in a different colore d ink) [large white rectangular label] / Cybocephalus californicus Horn Det: Trevor Smith (printed) [white rectangular label].

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54 Other material examined. CANADA: BRITISH COLUMBIA: R. D. OkanaganSimilkameen, Penticton, Dog Lake, Se ptember 23, 1927, coll. W. Mathers (3 1 OSAC; 1 1 TRSC); R. D. Okanagan-Similkameen, Summerland, August 15, 1955, coll. M. D. Proverbs, feeding on immature Phenacaspis pinifolia (4 3 CNCI; 1 1 TRSC); R. D. Okanagan-Similkameen, Summerland, August 21, 1964, Pinus ponderosa (1 CNCI); R. D. East Kootenay, Wardner, August 21, 1977, coll. B. F. & J. L. Carr (1 CNCI); UNITED STATES: ARIZONA: Apache Co., Adamana, May 7, 1903, coll. H. S. Barber (2 USNM); Apache Co., Canyon de Chelly, May 30, 1974, coll. K. Stephan (1 LSAM); Apache Co., Jct. I-40 & Hwy. 191, August 20-22, 1999, coll. E. Riley & M. Yoder (1 TAMU); Cochise Co., Dragoon Mts. Wood Canyon, April 29, 1972, coll. K. Stephan (1 FSCA); Cochise Co., Cochise Stronghold Recreation Area, August 3, 1990, coll. B. F. & J. L. Carr (1 CNCI); Coconino Co., Fredonia, August 9, 1966, coll. B. F. & J. L. Carr (1 1 CNCI); Coconino Co., Walnut (1 SEMC); Coconino Co., Walnut, coll. Wickam (1 SEMC); Coconino Co., Walnut, July, 1922, coll. Wickam (1 SEMC); Coconino Co., Williams, May 26, coll. Barber & Schwarz (1 USNM); Gila Co., Rooseve lt Reservoir, September 29, 1980, coll. B. F. & J. L. Carr (1 CNCI); Graham Co., S. Graham Mts. 5000ft, July 20, 1974, coll. K. Stephan (1 LSAM; 3 ,1 FSCA); Maricopa Co., Apache Lake, September 23, 1989, coll. B. F. & J. L. Carr (1 CNCI); Maricopa Co., Phoenix, January 24, 1930, coll. S. Flanders (1 UCRC); Mohave Co., Hot Springs, June 25, coll. Barber & Schwarz (2 USNM); Navajo Co., Winslow, July 31, coll. Barber & Schwarz (2 USNM); Pima Co., Greaterville, October 8, 1980, coll. B. F. & J. L. Carr (1 CNCI); Pima Co., Vail, May 3, 1975, coll. K. Stephan (1 FSCA); Pima Co., Tucson,

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55 December 21, coll. Hubbard & Schwarz (2 3 USNM); Pima, Co., Tucson, January 5, coll. Hubbard & Schwarz, Prosopis juliflora (1 USNM); Pinal Co., Oracle, January 9, June 29, July 5, 6, coll. Hubbard & Schwarz (3 8 USNM); Pinal Co., Superior, September 22, 1989, coll. B. F. & J. L. Carr (1 CNCI); Santa Cruz Co., Santa Rita Mts., 5000 to 8000 ft., July, coll. F. H. Snow (1 SEMC); Santa Cruz Co., Santa Rita Mts., June 20, coll. Hubbard and Schwarz (1 USNM); Yuma Co., Ft. Yuma, January 21, coll. Hubbard & Schwarz (1 USNM); CALIFORNIA: Butte Co., Chico, May 14, 1942, coll. E. H. Fosen (5 1 USNM; 1 1 TRSC); Colusa Co., Letts Lake, April 30, 1980, coll. Fred G. Andrews, S. Kuba, & T. D. Eichlin (1 1 CSCA); Contra Costa Co. Mt. Diablo, Ap ril 5, 1952, coll. R. Schuster (3 5 EMEC; 1 1 TRSC); Contra Costa Co., 3mi S. Martin ez, November 26, 1987, coll. L. G. Varela, on pear foliage (27 16 EMEC; 3 4 TRSC; 1 3 FSCA); Contra Costa Co., Albany, reared in Albany insectary (2 EMEC); Fresno Co., Selma, May 11, 1950, coll. P. DeBach, on red scale on grapefruit (1 UCRC); Fresno Co., Fresno, October 16, 1951 (2 EMEC); Imperial Co., Brawley, Apr. 13, 1976, ex. Lemon trees (1 1 CSCA); Imperial Co., El Ce ntro, May 4, 1951, coll. B. H. Harrigan, on red scale (1 UCDC); Imperial Co., Brawley, February 26, 1959, coll. E. I. Schlinger, collected by vacuum insect net in alfalfa field (1 UCRC); Imperial Co., Brawley, January 29, 1959, coll. E. I. Schlinger, Medicago sativa (1 UCRC); Imperial Co., Imperial Valley, June 11, 1971, coll. Flanders, on citrus (1 1 UCRC); Inyo Co., Bishop, July 26, 1921, coll. L. L. Muchmore, sage (4 3 LACM; 1 TRSC); Inyo Co., Westgard Pass, August 20, 1960, coll. E. I. Schlinger, on Chrysothamnus (2 2 UCRC; 1 TRSC); Kern Co., 12mi. NW Rosamond, July 22, 1955, coll. R. A. Flock, Lycium cooperi A.

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56 Gray (1 UCRC); Kern Co., 7mi. NW Mohave, July 16, 1965, coll. C. W. OBrien, ex. Lycium cooperi (3 5 EMEC; 4 4 FSCA; 1 1 TRSC); Kern Co., Ft. Tejon, July 3, August 3, 1930 (9 8 TAMU); Los Angeles Co., July, coll. Coquillett (1 USNM); Placer Co., August, coll. A. Koebele (1 USNM); Placer Co., Towle, Mar. 7, 1913, ex. Aulacaspis manzanitae (1 CSCA); Plumas Co., Lake Almanor, Apr. 23, 1971, coll. Fred G. Andrew s, ex. pine litter (1 CSCA); Los Angeles Co., Pomona (1 1 LACM; 2 INHS); Los Angeles Co., Arroyo Seco, February 21, 1971, coll. A Mayor (1 UCRC); Los Angeles Co., Little Rock, September 3, 1944, April 8, 1945, coll. G. P. Mackenzie (3 ,UCRC); Los Angeles Co., Los Angeles, February 1944, coll. R. H. Smith (4 UCRC; 1 TRSC; 1 FSCA); Los Angeles Co., Vincent, August 4, 1952, coll. Timberlake, on Atriplex canescens (1 UCRC); Merced Co., Merced, July 18, 1949, coll. R. L. Doutt, ex. Lepidosaphes ficus (1 EMEC); Monterey Co., UC Big Creek Reserve, Big Devils Ck. conf luence, 36.077 N 121.594 W, May 26-27, 2002, coll. S. Lew (1 SBMN); Orange Co., Santa Ana, December 18, 1936, coll. C. E. Norland, Diaspis echinocacti (3 4 LACM; 1 1 TRSC); Orange Co., Costa Mesa, January 21, 1943, (rest illegible) (1 UCRC); Placer Co., February 7, 1913, coll. B. B. Whitney (1 2 UCRC); Riverside Co., Mecca, February 25, 1914, coll. J. D. Neils, feeding on Parlatoria blanchardi (1 USNM); Riverside Co., Indio, May, 1926, coll. F. S. Stickney, ea ting Marlatt scale (6 6 USNM); Riverside Co., Mecca, April 17, 1967, coll. B. F. & J. L. Carr (1 3 CNCI); Riverside Co., Ripley, June 25, 1946, coll. W. F. Barr, Pluchea sericea (1 WFBM); Riverside Co., Riverside, March 29, 1989, coll. F. D. Bennett, cactus (4 FSCA); Riverside Co. Oasis, July 1967, coll. Fred G. Andrews (1 CSCA); Riverside Co., 31mi N Blythe, Apr. 27, 1978, coll. A. R.

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57 Hardy and Fred G. Andrews, collected on Pluchea sericera (1 CSCA); Riverside Co., 5 mi. N Aguanga, 116 52 30 W 33 32 30 N, August 17, 1978, coll. J. B. Woolley (2 UCRC); Riverside Co., Riverside, August 23, 1965, coll. M. E. Irwin (2 UCRC); Riverside Co., Mecca, November 17, 1970, coll. R. C. Dickson, on sticky board (2 2 UCRC; 1 TRSC; 1 FSCA); Riverside Co., Riverside, August 1, 1924, feeding on Aleyrodids, (rest is illegible) (2 UCRC; 1 1 TRSC); Riverside Co., Riverside, March 1926, on Aphidiotus on walnut, (rest is illegible) (1 2 UCRC); Riverside Co., Riverside, January 20, 1927, coll. Timberlake (rest illegible) (1 UCRC); Riverside Co., Coachella Valley, February 23, 2000, coll. G. R. Ballmer, on lettuce (1 UCRC); Riverside Co., Whitewate r Canyon, 650m, 33 57 18 N 116 38 39 W, September 4, 1999, coll. Yanega and Gates (1 UCRC); Riverside Co., Indio, 1mi. E. Jefferson Street, Kennedy Ranch, Ju ly 1963, coll. H. T. Reynolds, ex. cotton suction machine (2 UCRC); San Bernardino Co. 2mi. W. Phelan, June 7, 1958, coll. E. I. Schlinger, ex. scale on manzanita (1 UCRC); San Bernardino, Morongo, September 20, 1944, on Acacia greggi (rest is illegible) (2 UCRC); San Diego Co., San Diego, 1922, coll. Armitage, on cactus (2 2 UCRC); San Luis Obispo Co., Cuyama River, nr. Cuyama, August 6, 1999, coll. G. R. Ballmer, on Atriplex canescens root grown (1 UCRC); San Luis Obispo Co., Carri zo Plain N. M., Caliente Ridge, 35 N 119 82 W, D ecember 5, 2003 January 1, January 1-24, February 29March 17, March 17-April 2, 2004, coll. M. Cate rino, malaise, flight intercept, unbaited pitfall (2 2 SBMN; 3 TRSC); San Luis Obispo Co., Carrizo Plain N. M., Selby Campground, 35 N 119 W, February 7-29, January 1-24, February 29March 17, 2004, coll. M. Caterino, flight intercept (1 2 SBMN); Santa Clara Co.,

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58 San Jose, June (1 OSUC); Santa Clara Co. (1 OSUC); Sonoma Co., Healdsburg, Apr. 21, 1971, ex. Prunus (1 CSCA); Tehama Co., Mill Creek, August 31, 1947, coll. Timberlake, (rest is illegible) (1 UCRC); Tulare Co., Visalia, April 21, 1930, collected from sugar pine (2 LACM); Yuba Co., coll. E. J. Branigan (2 2 UCRC; 1 1 TRSC); COLORADO: Las Animas Co., June 16, 1982, coll. B. F. & J. L. Carr (1 CNCI); IDAHO: Boise Co., 1mi. S Ten Mile, June 14, 1988, coll. A. Allen, sweeping (1 AAIC); Canyon Co., Parma, 2224 ft., June 15, 1929, coll. C. Wakeland (1 WFBM); Canyon Co., 4.8 NW Walters Ferry, May 16, 1995, coll. W. F. Barr, sweeping Atriplex canescens & confertifolia (1 WFBM); Elmore Co., Mt. Home, 3138 ft., July 31, 1952, coll. W. F. Barr, Artemisia (1 OSAC); Fremont Co., Warm River Camp Campground, June 7, 1986, coll. B. F. & J. L. Carr (1 CNCI); Latah Co., Moscow, April 17, 1994, coll. M. M. Furniss (1 ,WFBM); Owyhee Co., Harpers Fairy Crossing, August 20, 1979, coll. A. Allen, on sage brush (1 AAIC); Twin Falls Co., Shoshone Falls, July 10, September 21, 1975, coll. A. Allen, sweeping sagebrush (3 AAIC); Twin Falls Co., August 5, 1978, coll. A. Allen, sweeping Artemisia (1 AAIC); Twin Falls Co., Derkies Lake, Snake River Canyon, June 29, 1977, coll. A. Allen, sweeping sagebrush (1 AAIC); MONTANA: Roosevelt Co., Snowden Bridge, June 11, 1991, coll. D. L. Gustafson (1 MTEC); Powder River Co., 5mi W. Broadus, June 7, 1999, coll. D. L. Gustafson (1 MTEC); Madison Co., 5mi E. Norris, Bear Trap Prim. Area Madison, 5000 ft., July 26, 1986, S. M. Fondriest (1 MTEC); NEVADA: Elco Co., Ruby Mtns. 7000 ft., May 14, 1975, coll. James H. Baker (4 USNM); NEW MEXICO: (1 1 INHS); Eddy Co., Lincoln National Forest, 4.5mi SW Queen, Hwy. 137, 1675m, 32 N 104 40 W, August 15-25, 2001, coll. J. C. Schaffner

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59 (1 TAMU); Socorro Co., Beartrap Canyon, Mt Withington, July 31, 1990, coll. B. F. & J. L. Carr (1 CNCI); OKLAHOMA: Latimer Co., March 1986, coll. K. Stephan (2 TAMU); OREGON: Klamath Co., Klamath Falls, June 6, 1956, coll. Joe Schuh (1 3 OSAC); Klamath Co., Upper Klamath Lk., Geary Canal, May 23, 1958, coll. Joe Schuh (1 OSAC); Wasco Co., Bear Springs, Oc tober 6, 1940, coll. K. M. & I. M. Fender (1 OSAC); TEXAS: Brewster Co., Big Bend Nat. Pk., 14mi E. Panther jct., September 9, 1985, coll. W. F. Barr, Croton (1 WFBM); Cameron Co., Brownsville, January 18, 1915, coll. Timberlake, assoc., with Dactylopius (1 1 UCRC); Gaines Co., Seminole, June 13, 1983, coll. B. F. & J. L. Carr (1 CNCI); Hudspeth Co., Indio Mountains Research Station, vic. Indio ranch house, 4040 ft., 30 N 105 W, June 12-13, 2002, coll. E. G. Riley, R. Diaz & M. J. Yoder (1 2 TAMU; 1 1 TRSC); Hudspeth Co., Indio Mountains Research Station, Cougar Canyon, 4040 ft., 30 N 105 W, Marc h 30-April 12, 2002, coll. A. R. Gillogly, Malaise Trap (1 TAMU); Hudspeth Co., Indio Mountai ns Research Station, Squaw Spring, 30 N 105 W, April 12-13, 2002, coll. E. G. Riley, & M. J. Yoder (1 TAMU); Hudspeth Co., Indio Mountains Research Station, Squaw Creek, 4200 ft., 30 N 105 W, Marc h 30-April 12, 2002, coll. R. Caesar & A. R. Gillogly (1 2 TAMU); Kennedy Co., 13.5mi S Sarita, October 11, 1994, coll. E. G. Riley (1 TAMU); Pecos Co., Hwy. 385 rest stop, 28mi S Ft. Stockton, April 19, 1997, coll. E. Riley (1 TAMU); Presidio Co., Big Bend Ranch St. Nat. Ar., Aqua Adento, June 1823, 1990, coll. D. Judd, Malaise Trap (1 7 TAMU; 1 1 TRSC); San Augustine Co., 10mi SE Broaddus, Coleman Ce metery, March 19, 1994, coll. E. Riley (1 TAMU); Travis Co., vic. Long Ho llow Ck., 30, May 8, 26, 1993,

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60 coll. Alexander, Quinn, Riley, Wharton, et al., on Juniperus ashei (3 4 TAMU; 1 1 TRSC); Travis Co., vic. Long Hollow Creek and Cypress Creek, 30, 3058, March 26, June 18-19, 1994, coll. M. Quinn, E. Riley, R.Wharton, on Quercus buckleyi (2 TAMU); Val Verde Co., Amistad Reservoir, Hwy 406, June 2, 2000, coll. E. G. Riley (3 4 TAMU; 1 TRSC); UTAH: Washington Co., St. George, coll. Wickham (1 SEMC); WASHINGTON: Benton Co., Hanford Works, 640 ft., July 30, 1952, September 17, 1952, September 24, 1952, coll. R. H. Whittaker, Sagebrush (3 OSAC). Remarks. Specimens are most often collected while sweeping brush, especially Artemisia spp. This beetle seems to occupy a similar niche to that of C. randalli, which is often collected in the same ar eas and on the same plants. While the distribution of this beetle is quite extensive, it does not seem to occur east of th e Mississippi River. Cybocephalus kathrynae T. R. Smith, New Species (Figs. 4-11, 4-12-4-16) Diagnosis. Male and female black. Antennal club has a smooth margin without a serrated edge and the scutellum has concave margins (Figs. 4-11), distinguishing this species from C. randalli, C. californicus, C. nigritulus and C. nipponicus Etymology. This species is named for Kath ryn Lang Zara-Smith, wife of the species names author. Description. Male. Form: Elongate oval; strongly convex dorsally. Length: 1.31.6 mm (measured from apex of clypeus to apex of elytra); width: 0.8-1 mm (measured at base of elytra). Color: Head, thorax, elytra and unders ide dark brown or black, the extreme edge of elytral apices with brown bor der. Front legs and antennae light brown or brown, middle and hind legs black or dark brown. Head: Broad and convex, clypeus

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61 wide and moderately produced. Eyes larg e and tear-shaped with internal margins distinct. Genae just visible from above and slightly explanate. Dorsal surface smooth but distinctly alutaceous with uniform punctati on. Antennae 11-segmented including a club with 3 antennomeres, club length about the size of eye. Club flat and distinctly separated from funicle and with a smooth ma rgin. First club antennomere wider than long, second club antennomere la rger than either first or third club antennomere and about as long as wide. Terminal club ante nnomere emarginate (Fig. 4-12). Antennomere 3 as long or longer than 4 and 5 combined. Pronotum: Strongly convex and shiny, lateral margin curved; anterior angle more narrowly arcuate than posterior. Surface evenly punctured and alutaceous. Scutellum: Alutacous and triangular with concave margins. Elytra: Uniform width narrowing at the apical 1 / 5 Strongly convex, sides almost parallel but slightly sinuous, apices rounded, length shorter than combined width (29:38). Very alutaceous along dorsal su rface and distinctly punctured, punctation ending before base of elytra, forming a na rrow impunctate area at base and sides. Median margin and edge of elytra bordered. Distinct striations at apices of elytra. Underside: Metasternum extremely alutaceous a nd roughly punctate, an d clothed in long coarse hairs. Abdominal ster nites alutaceous and punctate wi th long coarse hairs thinly covering the surface. Legs: All femora glossy with short hairs. Profemora narrowing slightly at basal end, mesoand metafemora extremely flattened and expanded. All tibiae slightly but distinctly curved and dilated toward the ap ex. Protibiae with short hairs along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides curving and convergent forming a sh arp point (Fig. 4-13). In profile,

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62 curved (Fig. 4-14). Median plate on surface very slightly elevated. Basal plate: Sides convergent and emarginate at apex (Fig. 4-15). Female. Nearly identical to male. Geographic distribution. Known only from the southern coast of Florida (Fig. 416). Hosts Cybocephalus kathrynae has been collected in very close proximity to Haliaspis nakaharai Howell and Haliaspis uniolae Takagi (Diaspididae). While these beetles have not actually been observed feedi ng on the aforementioned scale species, they were the only scale species in the beetles ha bitat and exhibited signs of predation. Type material examined The holotype, deposited in the MCZC, is a male specimen glued to a point with the following labels: USA: Florida, Monroe Co., Bahia Honda State Park southeast end of isla nd N24-W81 (printed) [white rectangular label] / V-14-2005, Trevor Smith & R. D. Cave, sifting sand dune leaf litter around Uniola paniculata (printed) [white rectan gular label] / HOLOTYPE Cybocephalus kathrynae T. R. Smith Det: Trevor Smith (pri nted) [red rectangular label]. The designated allotype, deposited in the MC ZC, is a female specimen glued to a point with the following labels: USA: FL: Monroe Co. Bahia Honda St. Pk. SE. end of island 3-III-2005, Trevor Smith, M. C. Thomas, sa nd litter, primary dune (printed) [white rectangular label] / ALLOTYPE Cybocephalus kathrynae T. R. Smith Det: Trevor Smith (printed) [blue rectangular label]. Designated paratypes are as follows: UNITED STATES: FLORIDA: Monroe Co., Bahia Honda State Pa rk, southeast end of island, December 1, 1999, coll: Paul Skelley, berl ese litter under tree s of 2 dunes (1 TRSC); Monroe Co., Bahia Honda State Park, south east end of island, N24-W81,

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63 May 14, 2005, coll: Trevor Smith & R. D. Ca ve, sifting sand dune leaf litter around Uniola paniculata (3 4 FSCA); Miami-Dade Co., Key Biscayne, Bill Baggs Cape Florida State Park., N25-W80, June 30, 2005, coll: Trevor Smith, sifting sand dune leaf litter (1 4 FSCA). Remarks. This species was collected by sifting sand and leaf litter in primary and secondary sand dunes where sea oats (Uniola paniculata L.) and seashore dropseed ( Sporobolus virginicus (L.)) were present. The sea oats were infested with H. uniolae and the seashore dropseed was infested with H. nakaharai It has only been collected on Bahia Honda Key in Monroe County and in Bill Baggs Cape Florida State Park in Miami-Dade County but may occur in other ar eas of Florida with naturally occurring sand dunes where sea oats are present and infested with armored scales. Cybocephalus nigritulus LeConte (Figs. 4-17-4-22) Cybocephalus nigritulus LeConte 1863: 64. Diagnosis. Male and female black and very glossy. Size similiar to C. californicus but slightly larger. Antennal club smaller than eye and truncate at terminal antennomere; each antennomere of club distinctly separated forming a serrated edge, distinguishing this species from C. kathrynae. The truncate terminal club antennomere distinguishes this species from C. randalli which has a rounded terminal antennomere. Protibia more dilated than in C. californicus and the translucent impunctate ar ea at the apices of elytra much smaller than in C. californicus. Male basal plate and median lobe are unique and easily distinguished from all other species. Redescription. Male. Form: Elongate, oval; contractile; strongly convex dorsally. Length: 1.0-1.55 mm (measured from apex of clypeus to apex of elytra);

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64 width: 0.85-1.19 mm (measured at base of elytra). Color: Head, thorax, elytra glossy black, underside black, with late ral margin of pronotum and apical margin of elytra yellowish and translucent. Front legs and antennae amber, middle legs amber or light brown, hind legs brown or dark brown. Head: Broad and convex, clypeus moderately produced. Eyes tear-drop shaped and large wi th internal margins distinct. Genae not visible from above but slightly explanate wh en viewed laterally. Dorsal surface smooth, but under high magnification very finely al utaceous, finely punctured. Antennae 11segmented including a club with 3 antennomeres; club length, the width of eye; club flat and distinctly separated from funicle and with a distinctly serra ted margin. First club antennomere wider than long, second club antenn omere much larger than first and third club antennomeres and about as long as wi de; terminal antennomere of club truncate, setose, and about as long as wide (Fig. 417). Antennomere 3 about the same length as antennomeres 4 and 5 combined. Pronotum: Strongly convex, under high magnification finely alutaceous with extreme lateral edges cu rving slightly and almo st straight, anterior angle more narrowly arcuate than posterior. Surface punctation uniform, but distinct, and somewhat sparse, surface shiny. Scutellum: Alutaceous and triangular with slightly convex margins. Elytra: Uniform width narrowing at the apical 1 / 5 Strongly convex, sides almost parallel but sli ghtly sinuous, apices rounded. Length somewhat shorter than combined width (34:44). Punctation distinct except along extreme edge of apex which is smooth and without punctation; under high magnification finely alutaceous. Median margin and apex of elytra bordered. Underside: Metasternum extremely alutaceous, roughly punctured and clothe d in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering surface. Legs: Femora glossy, broad,

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65 flattened and sparsely covered with short hairs. Proand mesofemora about same width from end to end. Metafemora expanded in mi ddle. Tibiae slightly but distinctly curved and dilated towards apex. Pro tibiae much more dilated than others and with short hairs along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or al most as long as 2 preceding tarsomeres combined. Median lobe: Sides parallel or slightly convergent curving into a large triangular point (Fig. 4-18, 4-19). In profile strongly curved from middle (Fig. 4-20). Median plate on surface elevated. Basal plate: Sides parallel sides at base, very slightly tapering towards apex. Apex flat with a dis tinct triangular protubera nce in center (Fig. 421). Female. Nearly identical to male. Geographic distribution. Alabama, Florida, Georgia, Indiana, Louisiana, Massachusetts, Michigan, Minnesota, Mississippi, Ontario, Pennsylva nia, Rhode Island, South Carolina (Fig. 4-22). Hosts. Like all cybocephalids, the primary f ood source of these beetles is armored scale insects (Diaspididae). Cybocephalus nigritulus has been seen in association with or reported feeding on C. pinifolia (Riley 1882), Fiorinia theae Green (Flanders 1934; from label data), Pseudaulacaspis cockerelli (Cooley) (from label data), and Pseudaulacaspis pentagona (Targioni-Tozzetti) (Collins and Whitcomb 1975). Type material examined The holotype in the MCZC is a male specimen glued to a point with the following labels: orange di sc / Type (printed) 6987 (handwritten) [dark orange rectangular label] / Cybocephalus nigritulus Lec. (han dwritten) [white rectangular label] / Cybocephalus nigritulus LeConte Det: Trevor Smith (p rinted) [white rectangular

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66 label]. The orange disc indicates that the sp ecimen was collected in the southern or gulf states. Two specimens in the MCZC that are possibly syntypes were also examined. One male and one female are glued to points on se parate pins, but have the same labels as follows: Detroit, Mich. (print ed) [white rectangular label] / Cybocephalus nigritulus LeConte Det: Trevor Smith [white rectangul ar label]. These specimens may be the Michigan specimens referred to by Horn (1879) in his redescripti on of this species. Other material examined. CANADA: ONTARIO: Kent Co., Tilbury, August 1967, coll. K. Stephan, sifting (1 FSCA); Essex Co., Wheatley, June 1967, coll. K. Stephan, coccids on aspen (1 FSCA); UNITED STATES: ALABAMA: Walker Co., Jasper, September 23, 1979, coll. Tim King, at light (1 AAIC); FLORIDA: Alachua Co., Gainesville, San Felasco Hammock St. Pres., Apr. 10, June 10, June 26, 2004, coll. Trevor Smith, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (20 20 TRSC; 8 10 FSCA); Alachua Co., Gainesville, University of Florida Natural Area., March 12, 2005, coll. Trevor Smith, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (20 20 FSCA); Alachua Co., Gaines ville, Haile Plantation, March 30, 2005, coll. Trevor Smith, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (5 5 FSCA); Alachua Co., Gainesville, Wilmont Gardens, March 8, 1974, coll. Fred Collins, on Camellia (1 USNM; 1 FSCA); Alachua Co., Gainesville, July 18, 1964, coll. R. E. White (1 USNM); Alachua Co., Gainesville, July 18,19, May 23, 24, 1964, coll. R. E.White (5 FSCA); Alachua Co., Gainesville, Devils Millhopper, September 28, 1997, co ll. Vince Golia, beating trees (1 FSCA); Alachua Co., Gainesville, May 28, 1988, coll. F. D. Bennett, pred. on Fiorina theae on Camellia (1 FSCA); Alachua Co., N. E. Gainesville, May 16, 2000, coll. J. Eric

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67 Cronin, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (1 1 FSCA); Alachua Co., Gainesville, San Fela sco Hammock St. Preserve, March 20, 2004, coll. M. C. Thomas, ex. Magnolia sp. (1 FSCA); Brevard Co., Titusville, SR 405, Enchanted Forest Sanct., White Trail, September 14-28, 2000, January 15-31, 2001, coll. Z. Prusak, P. J. Russell, S. M. Fulle rton, malaise trap, xeric oak hammock (1 1 UCFC); Dade Co., Miami, Snapper Creek Pl aza, October 20, 1991, coll. F. D. Bennett, Pseudaulacaspis cockerelli / Strelitzia ( 1 FSCA); Gadsen Co. Quincy, Hyw. 267, March 6, 1974, coll. Fred Collins, on Camellia (1 USNM); Gadsen Co., March 6, 1974, sawdust, Melia azedarach (1 FSCA); Orange Co., Orlando, UCF, April 5, 1999, coll. P. Russell, S. Fullerton, malais e trap, pond pine community/dahoon holly (2 UCFC); Orange Co., Orlando, UCF, April 19, 19 99, coll. P. Russell, S. Fullerton, malaise trap, maidencane marsh (2 UCFC); Orange Co., Orla ndo, UCF, May 11, 17, 24, June 2, 1999, coll. P. Russell, S. Fullerton, malaise trap, cypress forest (1 3 UCFC); Orange Co., Orlando, UCF, July 2, 1997, coll. S. Fullerton, malaise trap, long leaf pine/sand pine/turkey oak (1 UCFC); GEORGIA: Berrien Co., Alapaha, May 6, 2004, coll. Trevor Smith, on Magnolia grandiflora feeding on Pseudaulacaspis cockerelli (2 1 TRSC); LOUISIANA: East Baton Rouge Parish, Baton Rouge, Bluebonnet Swamp, August 18, 2000, coll. A. R. Cline, mv light (1 LSAM); MASSACHUSETTS: Suffolk Co., Boston, Arnold Arboretum, July 14, 1921, coll. Harold Morrison, swept from 5-leaf pines behind lab (1 USNM); MISSISSIPPI: Smith Co., July 22, 1956 (1 TRSC); PENNSYLVANIA: Allegheny Co. (1 SEMC); SOUTH CAROLINA: Dorchester Co., Summer ville, April 2, 1909, coll. J. G. Sanders, on tea scales (1 USNM).

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68 Remarks. These beetles are usually collected by hand, either beating or simply by gleaning and aspirating. They are also occasio nally picked up in Malaise traps. While they are sometimes found in urban areas, they seem to be collected more often in forested areas. The senior author collected hundreds of these beetles by spotting and aspirating them from Magnolia grandiflora L. infested with P. cockerelli While this beetle seems to be very widespread, ranging from Canada to Florida and west to the Mississippi River, there are relatively few specimens in collections. There is also relatively less known about the hosts of this particular species. Cybocephalus nipponicus Endrdy-Younga (Figs. 4-10, 4-23-4-27, 4-29) Cybocephalus nipponicus Endrdy-Younga 1971a: 244-245. Diagnosis. Male bicolored with yellow or tan head, proand mesosternum, antennae and legs and remainder of the body bl ack, thus distinguishing this species from all other Cybocephalus in North America. The female is almost completely black with yellow front legs and antennae, with remaini ng legs either light brown or brown. The antennal club is smaller than the eye, tr uncate and each antennomere is distinctly separated to form a serrated edge. The male basal plate and median lobe are distinctly different from other species. Redescription. Male. Form: Elongate, oval; contractile; strongly convex dorsally. Length: 1.0-1.35 mm (measured from apex of clypeus to apex of elytra); width: 0.9-1.30 mm (measured at base of elytra). Color: Head, prothorax, mesothorax, antennae and legs yellow or ye llowish brown; metathorax, elytra and abdomen black. Head: Broad and convex, clypeus moderately produ ced and comparatively narrow. Eyes large, oblong, with internal margins distin ct. Genae not visible from above and not

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69 explanate. Dorsal surface smooth, but under high magnification finely alutaceous, punctation fine. Antennae 11-segmented including a club with 3 antennomeres; club length short, about the length of eye. Club flat and distinctly separa ted from funicle and with a distinctly serrated margin. All club antennomeres wider than long and creating a serrated edge on inside margin of club. First club antennomere wider than long, second club antennomere larger than fi rst and third club ante nnomeres and slightly wider than long. Terminal club antennomere truncate, setose, and slightly wider than long (Fig. 4-23). Third antennomere slightly shorter than segments 4 and 5 combined. Pronotum: Strongly convex, lateral margins slightly curved or nearly straight; anterior angle more narrowly arcuate than posterior. Dorsal surface under high magnification finely alutaceous with fine punctation. Scutellum: Alutaceous and triangular with slightly convex margins. Elytra: Uniform width narrowing at the apical 1 / 5 Strongly convex, sides almost parallel but sli ghtly sinuous, apices rounded. Length somewhat shorter than combined width (33:40). Fine punctation at the base disappearing completely at distal edge but toward middle clearly three-armed a nd finely alutaceous. Median margin and apex of elytra bordered. Underside: Metasternum extremely alutaceous, roughly punctured and clothed in long coarse hairs. Abdominal sternites al utaceous and punctate with long coarse hairs thinly covering surface. Legs: Femora glossy, broad, flattened and sparsely covered with short hairs. Proand mesofemora about same width from end to end. Metafemora expanded in middle. Tibiae slightly but distinctly curved and dilated towards apex. Protibiae much more dilated than others a nd with short hairs along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as l ong as 2 preceding tarsomeres combined. Median

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70 lobe: Sides parallel at base, exte nding to a large apical triangle (Fig. 4-24), in profile strongly curved at tip (Fig. 4-25). Basal plate: Sides parallel at base, tapering apically (Fig. 4-26). Female. Similar to male except all black, late ral margin of the prothorax yellow or translucent and apical margin of elytra and middle and hind legs light brown or brown. Geographic distribution. Connecticut, Delaware, Florida, Hawaii, Maryland, Massachusetts, New Jersey, New York, Penns ylvania, Rhode Island, Texas, Virginia, Washington D.C. (Fig. 4-27), Asia, Europe, Micronesia, West Indies and South Africa. This is a recently introduced species nativ e to Southeast Asia, therefore specimens are rarely found in North American collections. For more information on the distribution of these beetles, see Drea and Carlson (1988), Jefferson et al. 1995, Van Driesche et al. (1998), Hudson et al. (2000), Deepak et al. (2003), Spichiger (2004), and HDA (2004). Hosts. This predator has been reported in association with or feeding on a large number of armored scale insects (Diaspididae): Aonidiella sp. (Endrdy-Younga 1971a), Aspidiotus destructor Signoret (Endrdy-Younga 1971a,b; from label data), Aulacaspis crawii (Cockerell) (from label data), Aulacaspis yasumatsui Takagi (Heu and Chun 2000; from label data), Chrysomphalus bifasciculotus Ferris (Hayashi 1978; Tanaka and Inoue 1980), Fiorinia externa Ferris (Spichiger 2004; from label data), Hemichionaspis sp. (Endrdy-Younga 1971a), P. cockerelli (Cooley) (from label data), P. pentagona (Targioni-Tozzetti) (Yasuda 1981; Endrdy-Younga 1971a,b), Diaspidiotus macroporanus (Takagi) (Tachikawa 1974), Quadraspidiotus perniciosus (Comstock) (Alvarez and Van Driesche 1998a), Unaspis euonymi (Comstock) (Alvarez and Van Driesche 1998b; Drea and Carlson 1988), and Unaspis yanonensis Kuwana (Tanaka and

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71 Inoue 1980). It has also been report ed feeding on the citrus red mite, P. citri (Tanaka and Inoue 1980). Material examined. UNITED STATES: CONNECTICUT: Hartford Co., Windsor, Connecticut Agricultural Experi ment Station, Valley Laboratory, Sept. 15, 2005, coll. R. S. Cowles, feeding on Fiorinia externa Ferris on Abies fraseri (Pursh.) (15 20 FSCA); FLORIDA: Collier Co., Naples, Apr. 25, 2002, coll. S. Krueger, on Cycas sp. (1 3 FSCA); Dade Co., Turnpike, Snapper Creek Plaza, June 25, 1990, coll. F. D. Bennett, ex. Pseudaulacaspis cockerelli on Ravenela madagascarensis (4 2 FSCA); Dade Co., S. Miami, Mar. 24, 2004, coll. P. Duetting, feeding on Aulacaspis yasumatsui on Cycas revoluta (10 10 TRSC); Dade Co., S. Miami, jct. US-1 & SW 141 St., March 24, Apr. 12, 2004 coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta (10 10 TRSC; 10 10 FSCA); Hillsborough Co., Plant City, May 14, 2004, coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta (10 10 TRSC; 10 10 FSCA); Hillsborough Co., Tampa, Downtown, July 14, 2005, coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta (10 10 FSCA); Manatee Co., Terra Ceia, Feb. 3, 1994, coll. M. Runnals, oleander (2 4 FSCA); Orange Co., Orlando, UCF, April 26, 19 99, coll. P. Russell, S. Fullerton, malaise trap, maidencane marsh (1, UCFC); Sarasota Co., Sarasota, 1315 38 th St., Jan. 8, 2005, coll. J. H. Frank, feeding on Aulacaspis yasumatsui on Cycas rumphii (3 3 TRSC); Seminole Co., Longwood, Oct. 7, 2004, coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta (7 5 TRSC); MARYLAND: Prince Georges Co., Beltsville, August 4, 1989, coll. R. Hendrickson, J. Drea, ex: Unaspis euonymi on: Euonymus sp. (3 2 TAMU; 1 1 TRSC); TEXAS: Brazos Co., Texas A&M

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72 West Campus, September 27, 2000, coll. M. Yoder (1 TAMU); INSECTARIES: CALIFORNIA: Orange Co., Anaheim, January 30, 1935, Insectary (3 3 LACM; 1 TRSC); Orange Co., Anaheim, October 15, 1952, coll. P. DeBach, on purple scale (1 5 UCRC); Orange Co., Santa Ana, Ju ly 1, 1944, (rest is illegible) (2 3 UCRC; 1 1 TRSC); Orange Co., Insectary, January, 1948 (1 UCRC);Ventura Co., Ventura, Rincon-Vitova In sectaries, Sept. 8, 2004 (10 10 TRSC; 10 10 FSCA); NEW YORK: Cayuga Co., Locke, IPM Laboratories Inc., June 2001, ex. China (1 6 FSCA; 1 1 LSAM); PENNSYLVANNIA: Delaware Co., Swarthmore, July 16-20, 1988, coll. Mike Rose, Texas A&M Quarantine Lab (native to Korea) (4 8 TAMU); Remarks. Cybocephalus nipponicus is the only U.S. cybocephalid which has a sexually dimorphic color pattern (Fig. 4-28). This species, now widely distributed throughout the eastern U.S., was misidentified as C. binotatus and released in south Florida in 1998 to combat A. yasumatsui (Anon. 1998; Howard and Weissling 1999) on sago palms ( Cycas spp.). These two species have been confused in the past as evidenced by Endrdy-Younga mixing specimens of C. binotatus (Fig. 4-28) and C. nipponicus (Fig. 4-29) in the description of C. binotatus in his 1968 monograph of the Palearctic cybocephalids. He later clarifie d this in his description of C. nipponicus and a redescription of C. binotatus (Endrdy-Younga 1971a). A male topotype of C binotatus was compared to the beetles released in Florida. Two black spots on the yellow pronotum, a metallic sheen on the elytra, and distinctive male genitalia clearly separate C. binotatus from C. nipponicus. It was believed that the 1998 release of C. nipponicus from Thailand was the first introduction of thes e beetles into the stat e of Florida (Howard

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73 et al. 1999). However, specimens dati ng back to 1989 were found in the FSCA and identified as C. nipponicus. Notes about type material of C. binotatus. The studied topot ype specimen in the BMNH is a male glued to a card with the following labels: W. Almora, Kumaon, H.G.C. (printed) [white rectangular label] / C ybocephalus binotatus, Grouv. (printed) [white rectangular label] / Ent. Mo. Mag. 1923 De t. G.C.C. (printed) [upside down white rectangular label] / H. G. Champion Coll. B. M. 1953-156 (printed) [white rectangular label] / Kirejtshuk det 1996 (printed) Cybo cephalus binotatus (handwritten) [white rectangular label]. This specimen is one of six collected by Champion (1923), of which one specimen of this series was designat ed as the neotype by Endrdy-Younga (1971a) when he could not find the original holotype deposited in the Assam Coll. (Grouvelle 1908). Cybocephalus randalli T. R. Smith, New Species (Figs.4-30-4-36) Diagnosis. Male and female black. Clypeus much broader and more extended than in C. californicus. Antennal club is unlike all other North American species in that it is as large or larger than the eye, and th e terminal antennomere of the club is rounded apically rather than truncate as in C. californicus, C. nigritulus, and C. nipponicus. Each club antennomere is distin ctly separated and forming a serrated edge, unlike C. kathrynae Front tibia is quite dilated, much more so than in C. californicus In males, the basal plate and median lobe are easily recognizable. Etymology. This species is named in honor of Randall Edwards Smith, father of the species names author.

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74 Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.2-1.3 mm (measured from apex of th e clypeus to apex of elytra); width: 0.850.95 mm (measured at base of elytra). Color: Head, thorax, elytra and underside black, lateral margin of pronotum and apical margin of elytra ye llowish and translucent; legs and antennae dark brown or black. Head: Broad and convex, clypeus wide and produced. Eyes comparatively small, oblong w ith internal margins distinct. Genae not quite visible from above and slightly e xplanate. Dorsal surface smooth, under high magnification distinctly alutaceous with fine punctation. Antennae 11-segmented including a club with 3 antennom eres; club large and about same size as or larger than eye. Club flat and distinctly separate from funicle and with a distin ctly serrated margin. First and second club antennomeres wider than long, terminal club antennomere longer than wide, apically rounded and quite setose (Fig. 4-30). Third antennomere shorter than antennomeres 4 and 5 combined. Pronotum: Strongly convex and glossy; anterior angle more narrowly arcuate than posterior. Surface finely punctate and alutaceous. Scutellum: Triangular with straight or slightly convex margins. Elytra: Uniform width narrowing at the apical 1 / 5 Strongly convex, sides almost parallel, apices rounded, length somewhat shorter than combined width (30:38 ). Very sparsely punc tate and distinctly alutaceous, punctation ending before the base of elytra, smooth at sides and base. Median margin and apex of elytra bordered. Apices of elytra with di stinct striations (Fig. 4-35). Underside: Metasternum extremely alutaceous and roughly punctate, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering the surface. Legs: All femora glossy, broad, flattened and sparsely covered with short hairs. Pr oand mesofemora about the sa me width from end to end.

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75 All tibiae slightly but distinctly curved a nd dialated toward the apex. Metafemora expanded in the middle. Protibiae much more dilated than the others and with short hairs along the outer margin. Mesoand metatibi ae with long stiff hairs along the outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres combined. Median lobe: Sides parallel before curving into a triangular apex (Fig. 4-31, 4-32). In profile, slightly curved at tip (Fig. 4-33). Median plate not elevated. Basal plate: Sides parallel at base, narrowing sl ightly before rounding at apex with a large concavity in the center (Fig. 4-34). Female. Nearly identical to male. Geographic distribution. California, Idaho, Nevada Utah, Washington (Fig. 436). Hosts. Specimens of C. randalli were collected while sweeping Artemisia spp., therefore it is a distinct possibility that these beet les feed on a scale found on Artemisia Type material examined The holotype, deposited in the MCZC, is a male specimen glued to a point with the following labels: USA: IDAHO FALLS CO. SHOSHONE FALLS 14 VII 1975 LEG: A ALLEN SWEEPING SAGE BRUSH (printed) [white rectangular label] / HOLOTYPE Cybocephalus randalli T. R. Smith Det: Trevor Smith (printed) [red rectangular label] The designated allotype, deposited in the MCZC, is a female specimen glued to a point with the following labels: Calif: Inyo Co. Saline Valley Dunes III-30-1976 cereal bowl pi t trap D. Giuliani (printed) [white rectangular label] / ALLOTYPE Cybocephalus randalli T. R. Smith Det: Trevor Smith (printed) [blue rectangular label]. Designated paratypes are as follows: UNITED STATES: CALIFORNIA: San Benito Co., 8.2 mi on Panoche rd. from I-5, Feb. 4, 1979

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76 Mar. 15, 1979, coll. A. J. Gilbert, Ethylen e glycol pit trap in Ephedra area (1 TRSC); San Benito Co., 8.2 mi on Panoche rd. from I-5, March 15 April 14, 1979, coll. A. J. Gilbert, antifreeze pit trap (1 CSCA); Riverside Co., Slover Ave. dump, 315m, 34 03 56 N 117 21 32 W, April 24, 2001, coll. Hawks and Yanega (1 UCRC); IDAHO: Twin Falls Co., Shoshone Falls, July 10, 1975, coll. A. D. Allen (1 USNM); Twin Falls Co., Shoshone Falls, September 21, 1975, coll. A. Allen, sweeping sagebrush (1 TRSC; 1 AAIC); NEVADA: Churchhill Co., 6mi East of Frenchman, August 22, 1972, coll. Stephen J. Chaplin (1 USNM; 1 TRSC); Nye Co., Mercury, July 21, 1964, on Aritri (1 USNM); Nye Co., Mercury, February 7, 1961 (1 USNM); UTAH: Washington Co., St. George July, coll. Wickham (1 CNCI); Washington Co., St. George (1 MTEC); WASHINGTON: Grant Co., Smyrna, June 19, 1932, coll. M. H. Hatch (1 FSCA); Remarks. This species is less frequently collected than the sympatric C. californicus Whether this beetle is actually less common than C. californicus or whether they just occupy a slightly different niche is unknown. While thes e beetles are usually picked up by sweeping, they occasionally show up in pitfall traps. This is probably a case of the optimal collecting technique not being employed to obtain this species.

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77 Figure 4-1. Complete Cybocephalus nipponicus genitalia: ED = ejaculatory duct; IS = internal sac; MA = muscle attachme nt; ML = median lobe; MS = median strut; Tbp = tegmen basal plate; Tdp = tegmen dorsal piece; TS = tegminal strut. Figure 4-2. Lateral habitus of Cybocephalus randalli.

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78 Figure 4-3. Truncate antenna of C. californicus Figure 4-4. Emarginate antenna of C. californicus. Figure 4-5. Median l obe, dorsal view, of C. californicus. Figure 4-6. Median lobe, dorsa l view (slide mounted), of C. californicus Figure 4-7. Median lobe lateral view, of C. californicus. Figure 4-8. Basal plate, ventral view, of C. californicus.

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79 Figure 49. U.S. states and Canadian provinces from which specimens of Cybocephalus californicus have been collected.

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80 Figure 4-10. Scutellum of Cybocephaus nipponicus Figure 4-11. Scutellum of Cybocephalus kathrynae.

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81 Figure 4-12. Antenna of C. kathrynae. Figure 4-13. Median lobe, dorsal vi ew (same as slide mounted), of C. kathrynae Figure 4-14. Median lobe, lateral view, of C. kathrynae Figure 4-15. Basal plate, ventral view, of C. kathrynae

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82 Figure 4-16. Collection localities of Cybocephalus kathrynae in Florida.

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83 Figure 4-17. Antenna of C. nigritulus Figure 4-18. Median lo be, dorsal view, of C. nigritulus. Figure 4-19. Median lobe, dorsa l view (slide mounted), of C. nigritulus Figure 4-20. Median lobe, lateral view, of C. nigritulus. Figure 4-21. Basal plate, ventral view, of C. nigritulus.

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84 Figure 4-22. U.S. states and Canadian provinces from which specimens of Cybocephalus nigritulus have been collected.

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85 Figure 4-23. Antenna of C. nipponicus. Figure 4-24. Median lobe, dorsal vi ew (same as slide mounted), of C. nipponicus. Figure 4-25. Median lobe, lateral view, of C. nipponicus Figure 4-26. Basal plate, ventral view, of C. nipponicus

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86 Figure 4-27. U.S. states from which specimens of Cybocephalus nipponicus have been collected.

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87 Figure 4-28. Dorsal habitus of C. binotatus Figure 4-29. Dorsal habitus of C. nipponicus.

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88 Figure 4-30. Antenna of C. randalli. Figure 4-31. Median lo be, dorsal view, of C. randalli Figure 4-32. Median lobe, dorsa l view (slide mounted), of C. randalli. Figure 4-33. Median lobe, lateral view, of C. randalli Figure 4-34. Basal plate, ventral view, of C. randalli

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89 Figure 4-35. Striations at the apices of elytra on C. randalli

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90 Figure 4-36. U.S. states from which specimens of Cybocephalus randalli have been collected.

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CHAPTER 5 THE CYBOCEPHALIDAE (COLEOPTER A) OF THE WEST INDIES AND TRINIDAD Introduction Sixteen species of Cyboce phalidae have been describe d from the New World, all belonging to the genera Cybocephalus Erichson and Pycnocephalus Sharp (Brethes 1922, Champion 1913, Reitter 1874, 1875, Sharp 1891, T.R. Smith 2006a, Waterhouse 1877). This is in stark contrast to the over 150 speci es belonging to six gene ra in the Old World. Smith and Cave (2006a) recently revised the five species known to occur in America north of Mexico, but the Neot ropical species have never been treated taxonomically. Cybocephalids are known to feed on a wide variety of hosts (Smith and Cave 2006a); however their most common food source is scale insects. In North America, Cybocephalus nipponicus Endrdy-Younga is a predator of Aulacaspis yasumatsui Takagi (Smith and Cave 2006b), Fiorinia externa Ferris (Spichiger 2004), and Unaspis euonymi (Comstock) (Drea and Carlson 1988, Alva rez and Van Driesche 1998a,b). In the western United States, Cybocephalus californicus Horn is a common natural predator of Ehrhornia cypressi (Ehrhorn) (Flanders 1934; Kartman 1946; Clausen 1940), Lecanium corni Bouch (Heintz 2001), and Diaspidiotus perniciosus (Comstock) (Heintz 2001). In the eastern United States, Chionaspis pinifoliae (Fitch) (Riley 1882), Fiorinia theae Green (Flanders 1934), Pseudaulacaspis cockerelli (Cooley) (Smith and Cave 2006a), and Pseudaulacaspis pentagona (Targioni-Tozzetti) (Collins and Whitcomb 91

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92 1975) are prey species of C. nigritulus LeConte. Pycnocephalus argentinus Brthes is a predator of Ceroplastes sp. (Brthes 1922, Parker 1951) in South America. To date, no cybocephalid species have been described from the West Indies. Due to the collecting techniques required to obtai n these beetles and th eir very small size, these beetles are often overl ooked and are rarely found in co llections. Cybocephalids do not come to light traps and are usually collected by beating vegetation, extracted from leaf litter samples, or from flight intercept traps. With collectors using collecting techniques such as sifting sand and leaf lit ter as well as the increased use of flight intercept traps in the West Indies, more of these beetles will be collected and subsequently described. The objectives of this study are to 1) present a key for the identification of the cybocephalids of the West Indies and Trinida d, 2) describe new West Indian species, and 3) record the distribution of each species in the Caribbean Basin. Materials and Methods Materials For this study, 222 specimens belonging to the genus Cybocephalus and 31 specimens belonging to the genus Pycnocephalus were examined. One female specimen from the Dominican Republic is at hand, but w ithout associated males their identification cannot be ascertained. Specimens were borrowed from the following institutions and private collections (name of the cura tor or owner in parentheses): AAIC Albert Allen Insect Collection, Boise, ID (Albert Allen) BMNH Natural History Museum, London [formerly British Museum (Natural History)], UK (Maxwell Barclay) FMNH Field Museum of Natural Hist ory, Chicago, IL (James H. Boone)

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93 FSCA Florida State Collecti on of Arthropods, Gainesville, FL (Paul Skelley and Michael Thomas) SEMC Snow Entomological Museum, University of Kansas, Lawrence, KS (Zack H. Falin) TAMU Texas A&M University, Department of Entomology, College Station, TX (Ed G. Riley) USNM United States National Museum, Smithsonian Institution, Washington D.C. (Gary Hevel) WIBF West Indian Beetle Fauna Proj ect Collection, Montan a State University, Bozeman, MT (Michael Ivie) Methods Genitalia were removed and disarticulat ion was carried out using the methods described in Smith and Cave (2006a). Definitions Median lobe: Also referred to as penis (Endrdy-Younga 1968, 1971a, 1971b; Kirejtshuk et al. 1997; Lupi 2003; Yu 1995a, 1995b). Basal plate: This is a reference to the basal plate of the tegmen (Endrdy-Younga 1968, 1971a, 1971b; Lupi 2003; Yu 1995a, 1995b). Key to the Cybocephalidae of th e West Indies and Trinidad 1. Head very short with extremely short, s lightly concave clypeus (Fig. 5-2); femora and tibiae of the posterior two pairs of legs extremely dilated and laminiform (Fig. 5-27); male with blue-green metallic sh een on head and apical half of pronotum.... ............................................... Pycnocephalus deyrollei (Reitter) new combination 1 Head not short, clypeus of normal size and not concave at the apex (Fig. 5-1); only hind femora dilated, middle femora a nd tibiae of the posterior two pairs of legs not or only slightly dilate d; male without metallic sheen.................................2 2(1) Antennal club without a serrated margin and terminal antennomere rounded (Fig. 5-11); scutellum with concave margins (Fig. 5-15)................................................... ................................................................................ Cybocephalus iviei new species 2. Antennal club with at least 2 antenn omeres forming a serrated margin and terminal segment truncate (Fig. 5-3); scutellum with straight or slightly convex margins (Fig. 5-16)..................................................................................................3

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94 3(2). Margin smooth between club ante nnomeres 1 and 2 but terminal club antennomere clearly much narrower than club antennomere 2 (Fig. 5-3); basal plate rounded (Fig. 5-6); median l obe as in Figs.5-4 and 5-5.................................... .Cybocephalus antilleus new species 3. Antennal club with a clearly serrated margin (Fig. 5-7)..........................................4 4(3). Male bicolored with h ead, prothorax, mesosternum and legs yellow or tan, rest of body black; basal plate coming to a rounded poi nt (Fig. 5-21); median lobe as in Figs. 5-19 and 5-20.............................. Cybocephalus nipponicus Endrdy-Younga 4. Male not bicolored...................................................................................................5 5. Male basal plate rounded with a slight concavity at the apex (Fig. 5-10); median lobe as in Figs. 5-8 and 5-9........................... Cybocephalus caribaeus new species 5(4) Male basal plate evenly rounded and without concavity (Fig. 5-25); median lobe as in Figs. 5-23 and 5-24........................ Cybocephalus geoffreysmithi new species Cybocephalus Erichson 1844 Cybocephalus Erichson 1844: 441-442. For a description of the genus, see Smith and Cave (2006a). Cybocephalus antilleus T. R. Smith, New Species (Figs. 5-3-5-6) Diagnosis. Male and female are black but w ith a matte-like appearance. The antennal club has a distinctive shape (Fig. 5-3) unlike any other species in the West Indies. In males, the basal plate (Fig. 56) and median lobe (Fig.5-4) are easily distinguished from thos e of all other species. Etymology. This species is named after the chai n of islands (the Antilles) in which it occurs. Description. Male. Form: Elongate round, contractile; strongly convex dorsally. Length: 1.3 mm (measured from apex of the cl ypeus to apex of el ytra); breadth: 0.7 mm (measured across elytral humeri). Color: Black, shiny but with a matte-like appearance.

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95 Head, thorax, and elytra black or dark brown, underside dark brown to black; legs and antennae dark brown. Head: Broad and convex, clypeus moderately produced and wide. Eyes large, round, facets distinct. Surface distinctly alutaceous and minutely but uniformly punctate. Clypeus wide and short. Genae not visible from above, slightly explanate when viewed laterally. Dorsal surface alutaceous and finely punctured. Antenna with 11 antennomeres including a 3segmented club. Club la rge, only slightly smaller than height of the eye, flat and distinctly separated from funicle. First and second club antennomeres wider than long, terminal club antennomere about as long as wide. Margin smooth between club antennomeres 1 and 2, but terminal club antennomere clearly much narrower than club antennomere 2 (Fig. 5-3). Antennomere 3 shorter than 4 and 5 combined. Pronotum: Alutaceous and punctuate. Strongly convex, lateral margins curved; anterior angles more narrowl y arcuate than posterior angles. Scutellum: Very small, alutaceous. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex, sides slightly sinuous and apices round, lengt h shorter than combin ed width (30:39). Punctation ending just before apices. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly puncture d, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctuate with long coarse hairs thinly covering the surface. Legs: All tibiae slightly but distinctly cu rved and dilated toward the apex. Protibiae with short hairs along the outer ma rgin. Mesoand metatibiae with long, stiff hairs along the outer margin. All femora glo ssy, wide, flattened and sparsely covered with short hairs. Proand mesofemora about the same width throughout their length. Metafemora expanded in the middle. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres. Median lobe: Sides parallel until converging sharply

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96 towards an acute, extended point at the apex (Fig. 5-4). In profile, slightly curved ventrally at tip (Fig. 5-5). Median pl ate only slightly elevated (Fig. 5-5). Basal Plate: Sides parallel at base, with a uniformly rounded tip (Fig. 5-6). Female. Similar to male except for genitalic structures. Distribution. Dominica. Material examined The holotype, deposited in the TAMU insect collection, is a partly disarticulated male specimen glue d to a point with the following labels: DOMINICA: St. Paul Par. Springfield Estate V-27-VI-12-1994 J. B. Woolley, 94/020 malaise trap (printed) [white rectangular label] / HOLOTYPE Cybocephalus antilleus T. R. Smith Det: Trevor Smith (printed) [red rect angular label]. The a llotype, deposited in the TAMU insect collection, is a female sp ecimen glued to a point with the following labels: DOMINICA: St. Paul Par. Springfield Estate V-27-VI-12-1994 J. B. Woolley, 94/020 malaise trap (printed) [white rectangular labe l] / ALLOTYPE Cybocephalus antilleus T. R. Smith Det: Trevor Smith (printed ) [blue rectangular label]. Paratype: Dominica: St. Paul Par., Pont Cass, 19VI-2004, coll. R. Turnbow (1 FSCA) Remarks. These beetles were collected in heavily forested habitat. Nothing is known about the preferred prey of these beetles. Cybocephalus caribaeus T. R. Smith, New Species (Figs. 5-7-5-10) Diagnosis. Male and female are black or dark brown. Each antennomere of the antennal club is distinctly separated to form a serrated edge and the terminal antennomere is truncate (Fig. 5-7). In males, the basal pl ate (Fig. 5-10) and median lobe (Fig. 5-8) are easily distinguished from all other species.

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97 Etymology. This species is named after the ge neral region (the Caribbean Sea) in which it occurs. Description. Male. Form: Round, contractile; strongly convex dorsally. Length: 1.35 mm (measured from apex of clypeus to apex elytra); breadth: 1.0 mm (measured across elytral humeri). Color: Head, thorax, elytra black or dark brown, underside dark brown to black; front legs light brown, middle and hind legs and antennae dark brown. Head: Broad and convex, clypeus moderate ly produced and relatively narrow. Eyes large, tear-shaped, facets distin ct. Dorsal surface distinctly alutaceous and minutely but uniformly punctate. Genae not visible from above and slightly explanate when viewed laterally. Antenna with 11 antennomeres including a 3-segmented club about the height of the eye. Club flat and distinctly separated from funicle. All club antennomeres wider than long; terminal club antennomere short and truncate (Fig. 5-7). Antennal club margin serrated. Antennomere 3 s horter than 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angles more narrowly arcuate than posterior angles. Alutaceous and punctuate. Scutellum: Triangular, alutaceous, and with margins slightly convex. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex, sides slightly sinuous and apices rounded, lengt h shorter than combined width (33:43). Punctation ending just before apices. Median margin and apices of elytra bordered. Underside: Abdominal sternites alut aceous and punctate with l ong coarse hairs thinly covering the surface. Metasternum alutace ous, roughly punctured, and clothed in long coarse hairs. Legs: All tibiae slightly but distinctly cu rved and dilated toward the apex. The protibiae with s hort hairs along the outer margin. Mesoand metatibiae with long, stiff hairs along the outer margin. All femora shiny, wide, flattened and sparsely covered

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98 with short hairs. Metafemora expanded in the middle. Proand mesofemora only very slightly dilated in the middle. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres. Median lobe: Sides extending parallel to one another, turning in towards the apex to form a sharp point, with a slight notch on either side of lobe of the way up towards the apex (Fig. 5-8). In profile, strongly curved from the middle (Fig 5-9). Median plate very elevated (Fig. 5-9). Basal Plate: Parallel sides at base narrowing slightly toward apex, emargi nate at the apical end (Fig. 5-10). Female. Similar to male except for genitalic structures. Distribution. Curaao Material examined The holotype, deposited in the US NM, is a disarticulated male specimen glued to a point with the following labels: CURAAO, Coral Specht 3 km. E. Willemstad 9-15 February 1987 W. E. Steine r & J. M. Swearingen (printed) [white rectangular label] / Flight-intercept yellow pa n trap in mesquite-acacia desert scrub near coast (printed) [white rectangular label] / HOLOTYPE Cybocephalus caribaeus T. R. Smith Det: Trevor Smith (printed) [red rectangu lar label]. The allot ype, deposited in the USNM, is a female specimen glued to a point with the following labels: CURAAO, Coral Specht 3 km. E. Willemstad 9-15 Februa ry 1987 W. E. Steiner & J. M. Swearingen (printed) [white rectangular label] / flight-i ntercept yellow pan tr ap in mesquite-acacia desert scrub near coast (printed) [w hite rectangular label] / ALLOTYPE Cybocephalus caribaeus T. R. Smith Det: Trevor Smith (p rinted) [blue rectangular label]. Remarks. This species was collected in desert habitat. Nothing is known about the diet or plant associa tions of this beetle.

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99 Cybocephalus iviei T. R. Smith, New Species (Figs. 5-11-5-15, 5-17) Diagnosis. Male and female are black. Size is extremely small (<1mm). The antennal club has a smooth margin (Fig. 5-11) without a serrated edge and the terminal segment is somewhat rounded. The scutellum has concave margins (Fig.5-15), distinguishing this species from all others in the West Indies. The apices of the elytra have a distinct invagination (Fig. 5-17), a character also unique to this species. Etymology. This species is named in honor of Dr. Michael Ivie, a thorough collector of microcoleoptera and th e first to collect this species. Description. Male. Form: Ovate; contractile; strongl y convex dorsally. Length: 0.80 mm (measured from apex of the clypeus to apex of elyt ra); breadth: 0.38 mm (measured across the elytral humeri). Color: Head, thorax, elytra black or dark brown, underside brown to dark brown; legs and antennae brown. Head: Broad and convex, clypeus moderately produced and wide. Eyes large, oblong with internal margins distinct. Genae extended laterally and easily visible from above and slightly explanate when viewed laterally. Dorsal surface alutaceous and finely punctured. Antenna with 11 antennomeres including a 3-segmented club only s lightly smaller than height of eye. Club flat and distinctly separa ted from funicle. First club antennomere wider than long. Second club antennomere larger than either first or third club antennomere and not as long as wide. Terminal club antennomere rounded and roughly trapezoidal in shape and about as long as wide (Fig. 5-11). An tennomere 3 slightly shorter than 4 and 5 combined. Pronotum: Strongly convex, lateral margins cu rved; anterior angles more narrowly arcuate than posterior angles. Surface distinctly punctured. Scutellum: Very wide with very concave margins (Fig. 5-15) ; alutaceous and very sparsely punctured.

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100 Elytra: Uniform width narrowing at the apical 1/5. Strongly convex dorsally, sides slightly sinuous and apices with a distinct i nvagination (Fig. 5-17). Length about half of combined width (17:30). Very alutaceous, distinctly punctured. Median margin and apices of elytra bordered. Underside: Abdominal sternites alut aceous and punctuate with long coarse hairs thinly covering the surface. Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Legs: All tibiae slightly but distinctly curved and dilated toward the apex. Protibiae with short hairs along the outer margin. Mesoand metatibiae with long, stiff hairs along the outer margin. All femora shiny, wide, flattened and sparsely covere d with short hairs. Proa nd mesofemora about the same width from end to end. Metafemora expanded in the middle. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres. Median lobe: Sides extending parallel to one another, turning in sharply towards the apex and coming to a blunt point (Fig. 5-12). In prof ile, strongly curved from middle (Fig. 5-13). Median plate only slightly elevated (Fig. 5-13). Basal Plate: Sides parallel at base, rounding to a slightly flat top (Fig. 5-14). Female. Similar to male except for genitalic structures. Distribution U. S. Virgin Islands : Buck Island (St. Croix), St. John, St. Thomas Type material examined The holotype, deposited in the USNM, is a male specimen glued to a point with the following la bels: Virgin Is: St. John Est. Concordia 12 May 1984, litter under cactus a nd agave, W. Muchmore (pri nted) [white rectangular label] / HOLOTYPE Cybocephalus iviei T. R. Smith Det: Trevor Smith (printed) [red rectangular label] / WIBF 014005 (printed with a barcode, upside down) [white rectangular label]. The allotype, deposited in the USNM, is a female specimen glued to a

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101 point with the following labels: Virgin Is : St. John Est. Concordia 12 May 1984, litter under cactus and agave, W. Muchmore (printed ) [white rectangular label] / ALLOTYPE Cybocephalus iviei T. R. Smith Det: Trevor Smith (printed) [blue rectangular label] / WIBF 013987 (printed with a bar code, upside down). Paratypes: UNITED STATES VIRGIN ISLANDS: BUCK ISLAND: Buck Island Reef National Monument, 140 ft., V-VI-1993, coll. Z. Hillis, fli ght intercept trap #14 (1 WIBF); Buck Island Reef National Monument, 340 ft., VII-VIII-1993, coll. Z. Hillis, flight intercept trap #15 (1 WIBF); Buck Island Reef National Monument, 340 ft., 30-III-29-VI-1995, coll. Z. Hillis and M. Hillis, flight intercept trap #15 (1 ,1 WIBF); Buck Island Reef National Monument, 9-V-1996, coll. Z. Hillis, M. Hillis and B. Phillips, flight intercept trap (1 ,2 WIBF) ; ST. JOHN: Maho Bay, 12-III-1984, coll. W. B. Muchmore, under trees nr. road (1 WIBF); Est. Concordia, 12-V-1984, coll. W. B. Muchmore, litter under cactus and dung (3 ,11 WIBF; 2 TRSC, 2, USNM); Est. Concordia, 12-V-1984, coll. W. B. Muchmore, litter under cactus and agave (5 ,10 WIBF; 1 ,1 MCZC; 1 ,3 FSCA); ST. THOMAS: Red Hook, 27-VII-1980, coll. M. A. Ivie, ex. Dead Stump (3 WIBF); Est. Nazareth, 40 ft., 26-VII-19-X-1994, coll. M. A. and L. L. Ivie, flight intercept #9 (2 2 WIBF). Remarks. While much smaller, this species has characteristics similar to C. kathrynae T. R. Smith which is found in southern Florida. Both species have similarly shaped antennal clubs and scutella. Cybocephalus iviei has almost always been collected in flight intercept traps or in l eaf litter. It was often collected in litter at the base of cacti and agave, therefore there is a good chance that one of their hosts is a scale insect found on one of these plants.

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102 Cybocephalus nipponicus Endrdy-Younga (Figs. 5-1, 5-16, 5-18-5-21) Cybocephalus nipponicus Endrdy-Younga 1971a: 244-245. Diagnosis. The male is bicolored with a yellow or tan head, proand mesosternum, antennae, and legs but the rema inder of the body is black, thus distinguishing this species from all other Cybocephalus in the West Indies. The female is almost completely black with yellow front legs and antennae, with the remaining legs either light brown or brown. The antennal club is smaller than the eye, truncate and each segment is distinctly separated to form a serrated edge (Fig. 518). The male basal plate (Fig. 5-21) and median lobe (Fig. 5-19-5-20) ar e distinctly different from t hose of all other species. Description. For descriptions of both sexes, see Smith and Cave (2006a). Distribution West Indies: Cayman Islands, St. Kitts / Nevis, Barbados; World: Asia, southern Europe, Micronesia, easte rn North America, and South Africa. Hosts. This predator has been reported feed ing on at least 14 species of armored scale around the world (see Smith and Cave ( 2006a) for references). According to label data, this beetle has only been recorded feeding on A. yasumatsui and Aspidiotus destructor Signoret in the West Indies. This be etle was introduced, using specimens collected in Florida, to Barbados where it quickly became established and contributes to the biological control of A. yasumatsui on ornamental king sagos, Cycas revoluta Thunberg (I. Gibbs, personal communication). Material examined. British West Indies: Cayman Islands: Grand Cayman, 15-VI-2001, coll. S. Frederic k, feeding on cycad scale on Cycas revoluta (8 6 FSCA); West Indies: St. Kitts / Nevis : St. Kitts, 14-II-1997, coll. R. D. Gautan, feeding on Aspidiotus destructor (6 3, BMNH); Nevis, nr. Charlestown, 9-XI-1995, coll. G.

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103 Watson, on coconut palm infested with whitefly (1 2, BMNH); Four Seasons Resort, 9-XI-1995, colls. G. Watson, M. Cock & E. Clarke, on coconut palm infested with Aspidiotus destructor and whiteflies (43 34, BMNH); Nisbet Plantation, 10-XI1995, coll. G. Watson, on coconut palm (20 14 BMNH); Nisbet Estate, 20-II-1997, coll. R. G. Booth, on coconut palm (3 2 BMNH); Potwork, 10-XI-1995, coll. E. Clarke & G. Watson, on coconut palm in fested with scale and whitefly (2 BMNH). Cybocephalus geoffreysmithi T. R. Smith, New Species (Figs. 5-22-5-25) Diagnosis. Male and female are black or dark brown. Each antennomere of the antennal club is distinctly separated to form a serrated edge and the terminal antennomere is truncate (Fig. 5-22). Scutellum with convex margins. In males, the basal plate (Fig. 525) and median lobe (Figs. 5-23, 5-24) are easily distinguis hed from all other species. Etymology. This species is named in honor of Geoffrey Edwards Smith, paternal grandfather of the species names author. Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.1 mm (measured from apex of clype us to apex of elytra); breadth: 0.9 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside dark brown, a narrow band along the lateral margin of pr onotum and posterior margin of elytra yellowish and translucent; front legs and antennae amber or light brown, middle and hind legs dark brown. Head: Broad and convex, clypeus moderately produced and narrow. Eyes large, oblong, with internal margins distinct. Genae not visible from above. Dorsal surface distinctly alutaceous and finely punc tate. Antennae 11-segmented including a club with 3 antennomeres, club length about size of the eye. Club flat and distinctly separated from funicle and with a distinctly serrated margin. First club antennomere

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104 slightly wider than long, second club antennomer e larger than either first or third club antennomere and wider than long. Terminal club antennomere truncate (Fig. 5-22), setose and wider than long. Antennomere 3 about as long as antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior. Surface di stinctly alutaceous with fine punctation. Scutellum: Very large, alutaceous and triangular with convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides s lightly sinuous and apices rounded. Length slightly shor ter than combined width ( 28:36). Uniformly punctate along dorsal surface, smooth at sides and base wi th a narrow impunctate area at apices of elytra. Appearing alutaceous under high magnification. Median margin and apices of elytra bordered. Apices w ith distinct striations. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdomin al sternites alutaceous and punctate with long coarse hair s thinly covering the surface. Legs: Femora glossy, broad, flattened and sparsely covered with short hairs. Proa nd mesofemora about the same width throughout length. All tibiae slightly but distinctly curved and dilated towards apex. The protibiae with s hort hairs along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres claw tarsomere as l ong or almost as long as 2 tarsomeres preceding it. Median lobe: Sides parallel or sl ightly divergent curving into a point (Fig. 5-23). In profile, strongly curved from mi ddle (Fig. 5-24). Median plate on surface elevated. Basal plate: Sides slightly convergent, evenly rounded at apex (Fig. 5-25). Female. Nearly identical to male. Distribution. Jamaica and Trinidad.

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105 Type material examined. The holotype, deposited in the BMNH, is a male specimen glued to a card with the followi ng labels: Jamaica: Hope River. 26.v.1908. Dr. M. Cameron. B.M. 1936-555 (printed) [white rect angular label with a yellow line running horizontally the length of the label] / British Mus. (printed) [white rectangular label] / HOLOTYPE Cybocephalus geoffreysmithi T. R. Smith Det: Trevor Smith (printed) [red rectangular label]. The designated allot ype, deposited in the BMNH is a female specimen glued to a card with the follo wing labels: Jamaica: Constant Spring. 29.vii.1908. Dr. M. Cameron. B.M. 1936-555 (print ed) [white rectangular label with a yellow line running horizontally the length of th e label] / British Mu s. (printed) [white rectangular label] / ALLOTYPE Cybocephalus geoffreysmithi T. R. Smith Det: Trevor Smith (printed) [blue rectangular label]. Paratypes: Jamaica: Rocky Spring, 8-III-1908, coll. M. Cameron (1 BMNH); Constant Spring, IV-1908, coll. M. Cameron (2 BMNH); Hope River, 26-V-1908, coll. M. Cameron (2 BMNH); Kingston, 16-II-1908, coll. M. Cameron (2 BMNH); Kingston, Palisadoes, 25-VIII-1966, coll. Howden & Becker (1 CNCI); Trinidad: St. Augustine, VI-1955, coll. F. D. Bennett, on Stachytarpheta sp. (2 2, BMNH) Remarks. Cybocephalus geoffreysmithi has been collected on Stachytarpheta therefore it stands to reason that its host is a scale found on this plant. Species of Stachytarpheta occur in both Jama ica and Trinidad. Pycnocephalus Sharp 1891 Pycnocephalus Sharp 1891: 373. Description. Form: Ovate and very convex; body contractile. Head: Very broad and short; deflexed (Fig. 5-2). Epistoma slightly prolonged at middle. Mandibles in repose resting against the meta sternum, acute at tip with a small tooth posteriorly.

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106 Maxillae with one lobe. Antennae shorter or as long as width of h ead, antennal club flat with three antennomeres, antennal grooves small and convergent. Scape greatly enlarged and of a particular shape (Fig. 5-26). Thorax: Pronotum margined at base, covering the base of elytra, sides very short. Prosternum acutely carinate in front, not prolonged behind the procoxae, procoxal cavities open behind. Mesosternum broad, oblique. Metasternum not or only slightly protuberant an d with very short and sparse hairs. Both the mesoand metasternum are deeply impre ssed for reception of middle and hind legs. Scutellum: Small and broad, triangular. Elytra: Covering or near ly covering tip of abdomen, apices curved. Abdomen: Five visible ventral plates (omitting the small male anal plate), and 5 abdominal spiracles. Legs: Femora and tibiae of the posterior two pairs of legs dilated and laminiform. Tibiae simple. Tarsi four-segmented, each tarsomere slightly dilated ve ntrally, second and third segments bilobed, claws simple. Median lobe: Heavily sclerotized and dorsoventrally compressed. Tegmen: Fused. Pycnocephalus deyrollei (Reitter), New Combination (Figs. 5-2, 5-26-5-30) Cybocephalus deyrollei Reitter 1875: 55-56. Diagnosis. The male is black with a blue-green metallic sheen on the head, anterior portion of the pronotum, and scape. The female is black and without the metallic sheen. Size is ca 2.5 mm. The extremely large and dilated middle and hind tibiae as well as the extremely short, broad head (Fig. 5-2) distinguish this species from all other cybocephalids in the West Indies. Description. Male. Form: Elongate, ovate; contractile ; strongly convex dorsally. Length: 2.4-2.5 mm (measured from apex of clypeus to apex of elytra); breadth: 1.751.85 mm (measured acro ss elytral humeri). Color: Head and apical portion of pronotum

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107 dark but with a blue or green metallic sh een, posterior portion of pronotum, elytra, and underside black, front legs light brown or am ber, middle and hind legs brown or dark brown, scape dark with a blue -green metallic sheen, rema ining 10 antennomeres brown or amber. Head: Broad, convex and very short, clype us extremely short and broad, with a slightly concave apical margin. Eyes very large, fairly round and oblong with internal margins distinct. Genae not visible from a bove. Dorsal surface alutaceous and finely punctate. Antenna with 11 antennomeres incl uding a 3-segmented club about 1/3-1/2 the height of the eye. Club flat, distinctly se parated from funicle, and with a distinctly serrated margin. First and second club an tennomeres wider than long, terminal club antennomere longer than wide, apically rounded and setose (F ig. 5-26). Third antennomere subequal to antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior angles. Surface finely punctuate. Scutellum: Triangular with margin s slightly convex and sinuous; alutaceous and very sparsely punctured. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex, si des almost parallel and apic es rounded, length shorter than combined width (50:72). Distinctly alut aceous and punctate, punctation extending to lateral edges and almost to apices. Median and lateral margins of elytra bordered. Underside: Metasternum alutaceous and roughly punctured, with short coarse hairs uniformly distributed. Abdominal sternites alutaceous and punctate with long, coarse hairs thinly covering the surface. Legs: Protibiae expanded towards the distal end and setose, profemora narrowing slightly at the distal end. Mesoa nd metatibiae with long, stiff hairs along the outer margin. Mesoand metafemora and tibiae very alutaceous, laminiform and greatly dilated (Fig. 5-27). F our tarsomeres, claw tarsomere as long or

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108 almost as long as 2 preceding tarsomeres combined. Median lobe: Sides rounded and curving to a rounded tip (Fig. 528). A patch of long hairs on either side of lobe about halfway between base and apex (Fig. 5-28). In profile, sinuous and slightly curved at tip (Fig. 5-29). Median plate not elevated. Basal Plate: Sides angling in before forming a flat top (Fig. 5-30). Female: Similar to male but head, pronotum, and first antennal segment black and without blue-green metallic sheen. Distribution. Trinidad and Tobago, Venezuela, and Brazil Material examined. West Indies: Trinidad and Tobago: Trinidad, St. George Co., Anma Asa Wright Nature Centre, 7-VII-1994, coll. C. Chaboo, beating vegetation (1 SEMC); Port-of-Spain, St. Clair, Sa vanna, 24-X-1918, coll. Harold Morrison (1 USNM); Arouca V, 1953, coll. NLH Krauss (1 USNM); 26-II-1956, coll. V. F. Hayes, on coffee berries (1 USNM); Talparo, 24-V-7-VI-1990, coll. H. L. Dozier (4 FSCA; 1 1 TRSC); Talparo, July 25-VII-1991, coll. H. L. Dozier (1 FSCA); Arima Valley, Simla, W. Beebe tropical rese arch station, 12-VII-1989, coll. H. L. Dozier (1 4 FSCA); St. Augustine, Mt. St. Benedict Abby, 7-VII-1996, coll. B. K. Dozier (2 1 FSCA); Arima Valley, road at mountain crest, 2-VII-1990, coll. H. L. Dozier (1 FSCA); N. range, Arima-Blanchisseuse Rd., mile 10, 11-V-1985, coll. C. W. & L. B. OBrien (1 FSCA); St. Augustine, IX-1962, co ll. F. D. Bennett, on croton (1 ,BMNH); 1905, coll. C. E. Bryant (5 1 BMNH); St. George Co., Anma, Asa Wright Nature Centre, 7-VII-1994, coll. C. Chaboo, ex. beating vegetation (1 SEMC); Morne Bleu, 2700, 25-VIII-1969, colls. H. & A. Howden (1 FMNH); Balandra Bay, 23-III-1922 (1 FMNH). Remarks. When Reitter described Cybocephalus deyrollei in

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109 1875 there was only one described genus in the family Cybocephalidae. It was not until 1891 that Sharp described a second genus, Pycnocephalus. Sharp recognized that the size and shape of the head as well as the dilated middle and hind tibiae justified the creation of a new genus. Cybocephalus deyrollei clearly exhibits the features described by Sharp; therefore we place it in Pycnocephalus The type specimens of Pycnocephalus metallicus Sharp, the genotype, were also examined and compared to C. deyrollei. The identification of C. deyrollei specimens was based on the published description since type specimens were not available for examination.

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110 Figure 5-1. Ventral habitus of C. nipponicus. Figure 5-2. Ventral habitus of P. deyrollei

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111 Figure 5-3. Antenna of C. antilleus. Figure 5-4. Median l obe, dorsal view, of C. antilleus Figure 5-5. Median lobe lateral view, of C. antilleus Figure 5-6. Basal plate, ventral view, of C. antilleus.

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112 Figure 5-7. Antenna of C. caribaeus Figure 5-8. Median l obe, dorsal view, of C. caribaeus. Figure 5-9. Median lobe lateral view, of C. caribaeus. Figure 5-10. Basal plate, ventral view, of C. caribaeus.

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113 Figure 5-11. Antenna of C. iviei Figure 5-12. Median lo be, dorsal view, of C. iviei Figure 5-13. Median lobe, lateral view, of C. iviei Figure 5-14. Basal plate, ventral view, of C. iviei

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114 Figure 5-15. Scutellum of C. iviei Figure 5-16. Scutellum of C. nipponicus.

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115 Figure 5-17. Invaginations at the elytral apices of C. iviei

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116 Figure 5-18. Antenna of C. nipponicus. Figure 5-19. Median lo be, dorsal view, of C. nipponicus Figure 5-20. Median lobe, lateral view, of C. nipponicus Figure 5-21. Basal plate, ventral view, of C. nipponicus

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117 Figure 5-22. Antenna of C. geoffreysmithi Figure 5-23. Median lo be, dorsal view, of C. geoffreysmithi. Figure 5-24. Median lobe, lateral view, of C. geoffreysmithi. Figure 5-25. Basal plate, ventral view, of C. geoffreysmithi.

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118 Figure 5-26. Antenna of P. deyrollei Figure 5-27. Hind leg of P. deyrollei Figure 5-28. Median lo be, dorsal view, of P. deyrollei Figure 5-29. Median lobe, lateral view, of P. deyrollei Figure 5-30. Basal plate, ventral view, of P. deyrollei

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CHAPTER 6 THE CYBOCEPHALIDAE (COLEOPTERA) OF MEXICO Introduction Cybocephalid beetles are mainly predat ors of scale insects (Vinson 1959, EndrdyYounga 1968, Alvarez and Van Dreische 1998a, b, Smith and Cave 2006b) but have also been known to feed on whiteflies (Chandra and Avasthy 1978, Kirejtshuk et al. 1997, Tian and Ramani 2003), and other arthr opod pests (Tanaka and Inoue 1980, EndrdyYounga 1982). Twenty-two species of Cybocep halidae have been described from the New World, all belonging to the genera Cybocephalus Erichson and Pycnocephalus Sharp. This is a very poorly studied family of beetles in the We stern Hemisphere and only recently has there been any type of taxonomic study of this group. Smith and Cave (2006a) recently revised the five species know n to occur in America north of Mexico, and also the Neotropical species occurring in the West Indies and Trinidad (Smith and Cave 2007). However the Mexican sp ecies have never been revised. The objectives of this study are to 1) develop techniques for the identification of the cybocephalids of Mexico, 2) describe ne w Mexican species, and 3) report the known distribution of each species known to occur in Mexico. Material and Methods Materials For this study, 103 specimens belonging to the genus Cybocephalus and 8 specimens belonging to the genus Pycnocephalus were examined. One female Cybocephalus specimen from Tampico is at hand, but without associated males its 119

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120 identification cannot be ascer tained. Specimens were borrowed from the following institutions and private collections (name of the curator or owner in parentheses): AAIC Albert Allen Insect Collection, Boise, ID, (Albert Allen) BMNH Natural History Museum, London [formerly British Museum (Natural History)], UK (Maxwell Barclay) CSCA California State Collection of Arth ropods, Sacramento, CA, (Chuck Bellamy) CNCI Canadian National Collection of Insects, Ottawa, Ontario, CANADA (Serge Laplante and Pat Bouchard) EMEC Essig Museum of Entomology, University of California, Berkeley, CA, (Cheryl Barr) FSCA Florida State Collecti on of Arthropods, Gainesville, FL (Paul Skelley and Michael Thomas) OSMN Orma J. Smith Museum of Natu ral History, Albertson College of Idaho, Caldwell, ID (William Clark) UCRC University of Californi a Riverside, Entomology Research Museum, Riverside, CA (Doug Yanega) USNM United States National Museum, Smithsonian Institute, Washington D.C. (Gary Hevel) Methods Genitalia were removed and disarticulat ion was carried out using the methods described in Smith and Cave (2006a). Definitions Median lobe: Also referred to as penis (Endrdy-Younga 1968, 1971a, 1971b; Kirejtshuk et al. 1997; Lupi 2003; Yu 1995a, 1995b). Basal plate: This is a reference to the basal plate of the tegmen (Endrdy-Younga 1968, 1971a, 1971b; Lupi 2003; Yu 1995a, 1995b, Smith and Cave 2006a, 2007).

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121 Key to the Cybocephalidae of Mexico 1. Head extremely short and broad with ve ry short antennae and slightly concave clypeus; femora and tibiae of the posterior two pairs of legs extremely dilated and laminiform; male with green metallic sheen on head and apical half of pronotum; size >1.5 mm.................Pycnocephalus metallicus Sharp 1 Head not short, clypeus of normal si ze and not concave at the apex; only hind femora dilated, middle femora and tibiae of the posterior two pairs of legs not or only slightly dilated; male wit hout metallic sheen; size <1.5 mm...........................2 2(1) Terminal antennomere of the antennal club rounded (Fig.6-25).............................3 2 Terminal antennomere of the antennal club truncate (Fig. 6-16) or slightly emarginate (Fig. 6-6, 6-7)........................................................................................4 3(2) Head and pronotum with a greasy app earance; antennal club small, at most only about half the size of the eye............................ Cybocephalus aciculatus Champion 3 Head and pronotum without greasy app earance; antennal club large, about the same size as eye; basal plate and me dian lobe as in Figs. 6-26-6-28........................ ........................................................................... Cybocephalus randalli T. R. Smith 4(2) Terminal antennomere of the antennal club longer than wide (Fig. 6-16); basal plate and median lobe as in Figs. 6-17-6-19................ Cybocephalus new species 3 4 Terminal antennomere of the antennal club wider than long..................................5 5(4) Male basal plate em arginate at the apex..................................................................6 5 Male basal plate not emarginate at the apex............................................................7 6(5) Terminal antennomere very short and wide (Fig. 6-2); basal plate with very shallow emargination at the apex (Fig. 65); when viewed laterally the median plate on the median lobe only slightly elevated (Fig. 6-4); median lobe as in Fig. 3 ...................................................................................... Cybocephalus new species 1 6 Terminal antennomere about as long as wide (Fig. 6-33); basal plate deeply emarginate at the apex (Fig. 6-36); when viewed laterally the median plate on the median lobe is extremely elevated (Fig. 6-35); median lobe as in Fig. 34................ ...................................................................................... Cybocephalus new species 4 7(5) Male basal plate protuberant at the apex (Figs. 6-15, 6-23)....................................8 7 Male basal plate not protuberant at the apex...........................................................9

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122 8(7) Sides of male basal plate slightly convergent towards apex, with distinct protuberance at the apex (F ig. 6-23); median lobe as in Figs. 6-21 and 6-22........... ..............................................................................Cybocephalus nigritulus LeConte 8 Sides of male basal plate parallel, with a gradual and weakly discernible protuberance at the apex (F ig. 6-15); median lobe as in Figs. 6-13 and 6-14........... ...................................................................................... Cybocephalus new species 2 9(7) Male basal plate evenly rounded at apex (Fig. 6-10); when viewed laterally the median plate on the median lobe is very elevated (Fig. 6-9); median lobe as in Fig. 6-8 .................................................................. Cybocephalus californicus Horn 9 Male basal plate with a wide, flat apex (Fig. 6-32); when viewed laterally the median plate on the median lobe is not el evated (Fig. 6-31); median lobe as in Fig. 6-30............................................................. Cybocephalus schwarzi Champion Cybocephalus Erichson 1844 Cybocephalus Erichson 1844: 441-442. For a description of the genus, see Smith and Cave (2006a). Cybocephalus aciculatus Champion (Figs. 6-1, 6-11) Cybocephalus aciculatus Champion 1913: 71. Diagnosis. Male and female are black but w ith a matte-like appearance. The terminal club antennomere is rounded (Fig. 61), separating this species from all other Mexican Cybocephalus except C. randalli. This species differs from C. randalli by the greasy sheen on the head and pronotum, larger eyes with many more ommatidia than C. randalli, and the antennal club smaller than the eye. Redescription. Female. Form: Elongate oval; contra ctile; strongly convex dorsally. Length: 1.2-1.3 mm (measured from apex of clypeus to apex of elytra); breadth: 0.85-0.90 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside black, lateral margin of pronotum and posterior margin of elytra yellowish and translucent; legs and antennae brown or black. Head: Broad and convex with a

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123 slightly greasy appearance, clypeus produced, and slightly reflexed. Eyes large, oblong, with internal margins distinct. Genae just vi sible from above and slightly explanate when viewed laterally. Dorsal surface smooth under 100 times magnif ication, distinctly alutaceous and finely punctate. Ante nnae 11-segmented including a club with 3 antennomeres, club length small, about size of the eye. Club flat and distinctly separated from funicle and with a distinctly serrated margin. First club antennomere wider than long, second club antennomere la rger than either first or third club antennomere and wider than long. Terminal club antennomere rounded (Fig. 6-1), setose and about as long as wide. Antennomere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior. Surf ace distinctly alutaceous with fine punctation and a greasy appearance. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides slightly sinuous and apices rounded. Lengt h slightly shorter th an combined width (32:38). Uniformly punctate along dorsal surf ace, smooth at sides and base with a small impunctate area at apices of elytra. Appearing very alutaceous under 100 times magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long, coarse ha irs. Abdominal sternites alutaceous and punctate with long, coar se hairs thinly c overing the surface. Legs: Femora glossy, broad, flattened, and sparsely covered with short hairs. Proand mesofemora about the same width throughout le ngth. All tibiae slightly but distinctly curved and dilated towards ap ex. Protibiae with short ha irs along outer margin. Meso

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124 and metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Male. Unknown. Geographic distribution. Federal District: Mexico City (Fig. 6-11). Type material examined. The lectotype, deposited in the BMNH, is a female specimen glued to a card with the follo wing labels: LECTO-TYPE (printed) [white circular label with a purple outer margin] / Mexico city. (printed ) Flohr (handwritten) [white rectangular label] / Tr. Ent. Soc. L. 1913 det. Champion (printed) [white rectangular label] / Cybocephalu s aciculatus Ch (handwritten) [white rectangular label] / LECTOTYPE Cybocephalus aciculatus Champion Det: Trevor Smith (printed) [red rectangular label]. While this specimen is a syntype, prior to this publication it was not a valid lectotype, despite the round lectotype label placed on the specimen, because this designation was never published. The de signated paralectotype is as follows: Mexico: Federal District, Mexico City, coll. J. Flohr (1 BMNH). Other material examined. Mexico: Tamaulipas, Tampico, 6-?-1912, coll. E. A. Schwarz (1 BMNH). Remarks. With only female type specimens it is very difficult to identify this species with any certainty. Cybocephalus New Species 1 (Figs. 6-2-6-5, 6-11) Diagnosis. Male is dark brown. Terminal cl ub antennomere slightly emarginate, separating this species from C. aciculatus and C. randalli and very distinctive by being much wider than long (Fig. 6-2), easi ly differentiating this species from Cybocephalus

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125 new species 3. In males, the basal plate (Fi g. 6-5) and median lobe (Figs. 6-3 and 6-4) are easily distinguished from all other species. Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.6 mm (measured from apex of clype us to apex of elytra); breadth: 0.85-1.20 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside dark brown, lateral margin of pronotum and poste rior margin of elytra yellowish and translucent; legs and antennae brown. Head: Broad and convex, clypeus moderately produced, narrow, and slightly reflexed. Eyes large, oblong, with internal margins distinct. Genae not visible from abov e. Dorsal surface smooth under 100 times magnification, distinctly alutaceous and finely punctate. Antennae 11-segmented including a club with 3 ante nnomeres, club length about size of the eye. Club flat and distinctly separated from funicle and w ith a distinctly serrated margin. First club antennomere wider than long, second club antenn omere larger than either first or third club antennomere and about as long as wide Terminal club antennomere truncate (Fig. 6-2), setose and wider than long. Antennom ere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than pos terior. Surface distinctly alutaceous with fine punctation. Scutellum: Very large, alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides slightly sinuous and apices rounded. Lengt h slightly shorter th an combined width (35:41). Uniformly punctate along dorsal surface, smooth at sides and base with a large impunctate area at apices of elytra. App earing alutaceous under high magnification. Median margin and apices of elytra borde red. Apices with distinct striations. Underside:

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126 Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering the surface. Legs: Femora glossy, broad, flattened, and sparsely covered with short hairs. Proand mesofemora about the same width throughout length. A ll tibiae slightly but distinctly curved and dilated towards apex. Pr otibiae with short hair s along outer margin. Mesoand metatibiae with long, stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides parallel and curving into a poi nt, apex shaped like a spearh ead (Fig. 6-3). In profile, slightly curved from middle (Fig. 6-4). Median plate on surface slightly elevated. Basal plate: Sides slightly rounded, apex with a very shallow concavity (Fig. 6-5). Female. External morphology identical to male. Geographic distribution. Sinaloa: nr. El Palmito (Fig. 6-11). Type material examined. The holotype, deposited in the CNCI, is a partially disarticulated male specimen glued to a car d with the following labels: 15 mi. W. El Palmito, Sin. MEX. VII-29-64 H. F. Howden (printed) [white rectangular label]. The allotype, deposited in th e CNCI, is a female specimen glued to a point with the following labels: 15 mi. W. El Palmito, Sin. MEX. V II-28-64 H. F. Howden (printed) [white rectangular label]. One female paratype is labelled as follows: Mexico: Sinaloa 8 mi. W E. Palmito, 7-VIII-1964, coll. H. F. Howden (CNCI). Cybocephalus californicus Horn (Figs. 6-6-6-11) Cybocephalus californicus Horn 1879: 320-321. For a redescription of the speci es, see Smith and Cave (2006a).

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127 Diagnosis. Male and female are black, brown, or aeneous. Antennal club is smaller than the eye and truncate or slightly emarginate at the apex (Figs. 6-6 and 6-7), separating this species from C. aciculatus and C. randalli The terminal club antennomere is wider than long, unlike Cybocephalus new species 3, but not as short as Cybocephalus new species 1. In males, the basa l plate (Fig. 6-10) and median lobe (Figs. 6-8 and 6-9) are easily distinguished from all other species. Geographic distribution. Baja California and northern Mexico (Fig. 6-11). Material examined. Mexico: Baja California Sur, Las Barracas, 2-V-1986, coll. P. DeBach, Ex. cactus scale on Opuntia cholla (3 5, UCRC); 50.3 km. SE Guerrero Negro, 15-I-1974, E. L. Sleeper (2 CSCA); Baja California Norte, 6 mi. N Guerrero Negro, 16-III-1981, F. Andrews & D. Faulkner, collected in flowers of Sphaeralcea axillaris (1 1, CSCA); 12.7 km. E El Rosario, 30N 115W, 180 m., 18VII-1991-28-V-1992, coll. W. H. Clark, E. M. Clark, P. E. Blom & D. M. Ward, ethylene glycol trap (1 OSMN); 12.7 km. E El Rosario, km. 68, 30N 115W, 180 m., 7-II-1984-2-IV-1985, coll. W. H. Clark & P. E. Blom, ethylene glycol trap (1 OSMN); Sonora, Hermosillo, 28-VIII-1954, co ll. R. Debach, Ex. Aonidiella aurantii (Maskell) (6 4, UCRC); Punta Chueca, 18-IV-1978, Ex. Jojoba (9 11 CSCA); Nogales, 3-VIII-1966, coll. Schwenke (1 USNM); Tamaulipas, 18 mi. SE Estacion Manuel, 1-XI-1982, coll. J. Huber (1 CSCA). Remarks. Label data indicate that this species feeds on the armored scale Aonidiella aurantii (Maskell) and a scale insect infesting Opuntia ( Cylindropuntia ) sp. For a detailed host list north of Me xico see Smith and Cave (2006a).

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128 Cybocephalus New Species 2 (Figs. 6-12-6-15, 6-24) Diagnosis. Male and female are black. The antennal club is smaller than the eye and the terminal club antennomere is truncate and wider than long (Fig. 6-12), separating this species from C. aciculatus C. randalli, and Cybocephalus new species 3. The terminal club antennomere is not as short as in Cybocephalus new species 1. In males, the basal plate (Fig. 6-15) and median lobe (F igs. 6-13 and 6-14) are easily distinguished from all other species. Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.4 mm (measured from apex of clype us to apex of elytra); breadth: 1.0 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside black, posterior margin of elytra yellowish and tr anslucent; legs and antennae brown or black. Head: Broad and convex, clypeus moderately produ ced, narrow, and slightly reflexed. Eyes large, oblong, with internal margins distinct. Genae not visible from above. Dorsal surface smooth under 100 times magnification, di stinctly alutaceous and finely punctate. Antennae 11-segmented including a club w ith 3 antennomeres, club length about size of the eye. Club flat and distinctly se parated from funicle a nd with a distinctly serrated margin. First club antennomere wi der than long, second club antennomere larger than either first or third club antennomere and about as long as wide. Terminal club antennomere truncate (Fig. 6-12), setose and a little wider than long. Antennomere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior. Surface distinctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5

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129 Strongly convex, sides slightly sinuous and ap ices rounded. Length slightly shorter than combined width (34:40). Uniformly punctate along dorsal surface, smooth at sides and base with an impunctate area at apices of elytra. Appearing alutaceous under 100x magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering the surface. Legs: Femora glossy, broad, flattened and sparsely covered with short hairs. Proand mesofemora about the same width throughout le ngth. All tibiae slightly but distinctly curved and dilated towards ap ex. Protibiae with short ha irs along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides parallel or slightly divergent, curving into a triangular point (Fig. 6-13) In profile, strongly curved from middle (Fig. 6-14). Medi an plate on surface elevated. Basal plate: Sides parallel at base, rounded at apex with a very slight protuberance in the ce nter (Fig. 6-15). Female. Unknown. Geographic distribution. Jalisco : Puerto Vallarta (Fig. 6-24). Type material examined. The holotype, deposited in the USNM, is a disarticulated male specimen glued to a car d with the following labels: MEXICO Jalisco Puerto Vallarta 25 Jan 1984 G. E. Bohart (printed) [white rectangular label]. Cybocephalus New Species 3 (Figs. 6-16-6-19, 6-24) Diagnosis. Male and female are black. The antennal club is smaller than the eye and the club antennomere is truncate and much longer than wide (Fig. 6-16), separating

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130 this species from all others in Mexico. In males, the basal plate (Fig. 6-19) and median lobe (Figs. 6-17 and 6-18) are easily distinguished from all other species. Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.1 mm (measured from apex of clype us to apex of elytra); breadth: 0.9 mm (measured across elytral humeri). Color: Head, thorax, elytra, and underside dark brown, posterior margin of elytra transl ucent; legs and antennae brown or black. Head: Broad and convex, clypeus moderately produced narrow, and slightly reflexed. Eyes large, oblong, with internal margins distinct Genae not visible from above. Dorsal surface smooth under 100 times magnification, di stinctly alutaceous and finely punctate. Antennae 11-segmented including a club w ith 3 antennomeres, club length about size of the eye. Club flat and distinctly se parated from funicle a nd with a distinctly serrated margin. First club antennomere wi der than long, second club antennomere about as long as wide. Terminal club antennomere truncate (Fig. 6-16), se tose and longer than wide. Antennomere 3 as long or slightly l onger than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curv ed; anterior angle more narrowly arcuate than posterior. Surface distin ctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides s lightly sinuous and apices rounded. Length slightly shor ter than combined width ( 32:37). Uniformly punctate along dorsal surface, smooth at sides and base wi th a large impunctate area at apices of elytra. Appearing alutaceous under 100 times magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long, coarse hairs

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131 thinly covering the surface. Legs: Femora glossy, broad, fla ttened and sparsely covered with short hairs. Proand mesofemora about the same width throughout length. All tibiae slightly but distinctly curved and dilated towards ape x. Protibiae with short hairs along outer margin. Mesoand metatibiae with long, stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almo st as long as 2 tarsomeres preceding it. Median lobe: Sides parallel curving abruptly into a rounded protuberance, with a patch of long hairs on either side of the lobe at the apical curv ature (Fig. 6-17). In profile, strongly curved from middle (Fig. 6-18) Median plate on surface elevated. Basal plate: Sides divergent, apex with very larg e triangular emargina tion (Fig. 6-19). Female. External morphology identical to male. Geographic distribution. Tamaulipas: Tampico (Fig. 6-24). Type Material Examined. The holotype, deposited in the USNM, is a disarticulated male specimen glued to a card with the following labels: Tampico Mex (printed) 6-12 (handwritten) [white rectangul ar label] / EA Schwar z Collector (printed) [white rectangular label]. The allotype, deposited in the USNM, is a female specimen glued to a point with the following labels : Tampico Mex (printed) 24-12 (handwritten) [white rectangular label] / EA Schwarz Collector (printed ) [white rectangular label]. Cybocephalus nigritulus LeConte (Figs. 6-20-6-24) Cybocephalus nigritulus LeConte 1863: 64. For a description of the species see Smith and Cave (2006a). Diagnosis. Male and female are black and very glossy. The antennal club is smaller than the eye and truncate at the term inal antennomere (Fig. 6-20), distinguishing this species from C. aciculatus and C. randalli The terminal club antennomere is wider

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132 than long, unlike Cybocephalus new species 3. In males, the basal plate (Fig. 6-23) and median lobe (Figs. 6-21 and 6-22) are eas ily distinguished from all other species. Geographic distribution. Durango, Durango; San Luis Potos, Ciudad del Maz (Fig. 6-24). Material examined. Mexico: San Luis Potos: Ciudad del Maz, 24-V-1996, coll. H. L. Dozier (1, 1, FSCA); Durango: Durango, 26-XI-1909, coll. F. C. Bishop (1 USNM). Remarks. This species is typically found east of the Mississippi River in the United States, however specimens have been collected west of the Mississippi along the gulf coast (Smith and Cave 2006a). Therefore, it is not surprising th at this species is found in Mexico. For a detailed host list north of Mexico see Smith and Cave (2006a). Cybocephalus randalli T. R. Smith (Figs. 6-25-6-28, 6-41) Cybocephalus randalli T. R. Smith 2006. For a description of the species see Smith and Cave (2006a). Diagnosis. Male and female are black. The ante nnal club is as large as or larger than the eye (Fig. 6-25), separating this species from all other Mexican Cybocephalus In males, the basal plate (Fig. 6-28) and median lobe (Figs. 6-26 and 6-27) are easily distinguished from all other species. Geographic distribution. Baja California (Fig. 6-41). Material examined. Mexico: Baja California Sur, 30 mi. S El Arco, 6-III-1977, coll. H. Marz ((1 FSCA); Baja California Norte, El Arco, 10-VIII-1980, coll. Bill Clark (1 AAIC); Valle Montevideo La Laguna wash, 18 km. W Bahia Los Angeles, 28N 113W, 380 m, January 3-I-12-V III-1982, ethylene glycol pitfall trap (1

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133 AAIC); Sierra Juarez, 4 mi. S El T opo, 10-III-1962, coll. E. L. Sleeper (1 CSCA); km. 147, 2 mi. SE Rancho Sonora, 29N 114 55W, 600 m., 16-VII-1991-27-V-1992, coll. W. H. Clark, E. M. Clark, P. E. Blom & D. M. Ward Jr., ethylene glycol pitfall trap (3 6, OSMN); Mesa Palmarito,29 54N 114W, 800 m., 27-VIII-1989-27-IV-1990, coll. W. H. Clark, M. H. Clark & P. E. Blom, ethylene glycol pitfall trap (1 4, OSMN); Mesa Palmarito, 29 47N 114W, 800 m., 27-X-1990, ethylene glycol pitfall trap (1 OSMN); 2 km. E. El Berrendo West, 30-III-2005, ungrazed out of the oasis, ethylene glycol pitfall trap (1 OSMN); 9 km. NW Rancho Santa Ynes, 29 46N 114W, 550 m., 4-I-1982-25-VI II-1982, coll. W. H. Clark & P. E. Blom, ethylene glycol pitfall trap (1 1 OSMN); 9 km. NW Rancho Santa Ynes, 29 46N 114W, 550 m., 22-VIII-1982, coll. W. H. Clark & P. E. Blom, uv-light trap (1 OSMN); Valle Montevideo, 18 km. W Bahia de Los Angeles, 28 55N 113W, 380 m., 14-VII-1991-13-V-1992, coll. W. H. Clark & P. E. Blom, ethylene glycol pitfall trap (1 1 OSMN); Valle Montevideo, La Laguna wash, 28 55N 113W, 380 m., 19III-1991-14-VII-1991, coll. W. H. Clark, M. H. Cl ark, C. J. Clark, K. D. Clark & J. E. Luther, ethylene glycol trap (2 3, OSMN); San Agustin, 29 56N 114W, 9-IV1985-9-III-1986, coll. W. H. Clark & P. E. Blom, ethylene glycol pitfall trap (4 OSMN); 4 mi. E San Agustin, 29 59N 114W, 17-I-2001-14-XII-2001, coll. W. H. Clark & D. M. Ward, ethylene glycol pitfall trap (2 OSMN); 4.3 km. NE Pozo Aleman, 28 14N 113W, 300 m., 15VI-1990-19-V-1992, W. H. Clark & P. E. Blom, ethylene glycol pitfall trap (1 OSMN); 12 km. NE El Arco, 28 18N 113W, 300 m., 24-IV-1990-15-VI-1990, W. H. Clark, M. H. Clark & D. Ward Jr., ethylene

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134 glycol pitfall trap (1 OSMN); 1 km. NE Santa Catarina (ranch), 29 44N 115W, 20-III-1991-3-VII-1991, ethylene glycol pitfall trap (1 OSMN). Remarks. Unlike all other species of Cybocephalus, this species is frequently collected in pitfall traps. Cybocephalus randalli is typically collected in desert habitats and their reduced eyes may indicate that th is species spends part of its time under the sand. This behavior has been seen in Cybocephalus kathrynae T. R. Smith which is collected by sifting loose sand near gra ss clumps in Florida. Cybocephalus schwarzi Champion (Figs. 6-29-6-32, 6-41) Cybocephalus schwarzi Champion 1913: 72. Diagnosis. Male and female are dark brown. Th e antennal club is smaller than the eye and its apex is truncate (Fig. 6-29), unlike C. aciculatus and C. randalli. The terminal antennomere is wider than long, unlike Cybocephalus new species 3, but not as short as Cybocephalus new species 1. In males, the basal plate (Fig. 6-32) and median lobe (Figs. 6-30 and 6-31) are easily distinguished from all other species. Redescription. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.3 mm (measured from apex of clypeus to apex of elytra); breadth: 0.9 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside dark brown, lateral margin of pronotum and poste rior margin of elytra yellowish and translucent; legs and antennae brown or black. Head: Broad and convex, clypeus moderately produced, narrow, and slightly re flexed. Eyes large, oblong, with internal margins distinct. Genae not visible from above. Dorsal surface smooth under 100 times magnification, distinctly alutaceous and finely punctate. Antennae 11-segmented including a club with 3 ante nnomeres, club length about size of the eye. Club flat

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135 and distinctly separated from funicle and w ith a distinctly serrated margin. First club antennomere wider than long, second club antenn omere larger than either first or third club antennomere and about as long as wide Terminal club antennomere truncate (Fig. 6-29), setose and slightly wide r than long or about as long as wide. Antennomere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; anterior angle more narrowly arcuate than posterior. Surface distinctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides slightly sinuous and ap ices rounded. Length slightly shorter than combined width (32:40). Uniformly punctate along dorsal surface, smooth at sides and base with a large impunctate area at apices of elytra. Appearing alutaceous under 100 times magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long, coarse hairs thinly covering the surface. Legs: Femora glossy, broad, flattened and sparsely covered with short hairs. Proand mesofemora about the same width throughout length. A ll tibiae slightly but distinctly curved and dilated towards apex. Pr otibiae with short hair s along outer margin. Mesoand metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides parallel or slightly convergent, curving abruptly into a sharp point (Fig. 6-30). In profile, curved from the lower middle region (Fig. 6-31). Median plate on surface elevated. Basal plate: Sides parallel at base, rounding to a flat apex (Fig. 6-32). Female. External morphology identical to male.

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136 Geographic distribution. Tamaulipas: Tampico (Fig. 6-41). Type material examined. The lectotype, deposited in the BMNH, is a male specimen glued to a card with the follo wing labels: LECTO-TYPE (printed) [white circular label with a purple outer margin] / Tampico (printed) 18-12 (handwritten) [white rectangular label] / E. A. Schw arz Collector (printed) [white re ctangular label] / U.S. Nat. Mus.1913-253 (printed) [upside down white rect angular label] / Cybocephalus schwarzi Ch. (handwritten) [white rect angular label] / LECTOTYPE Cybocephalus schwarzi Champion Det: Trevor Smith (printed) [red rect angular label]. While this specimen is a syntype, prior to this public ation it was not a valid lectotype, despite the round lectotype label placed on the specimen, because th is designation was never published. A paralectotype is labelled as follows: Mexico: Tamaulipas, Tampico, 14-?-1912, coll. E. A. Schwarz (1 BMNH). Cybocephalus New Species 4 (Figs. 6-33-6-36, 6-41) Diagnosis. Male and female are dark brown. Th e antennal club is smaller than the eye and the terminal antennomere is truncate (Fig. 6-33), unlike C. aciculatus and C. randalli, and wider than long, unlike Cybocephalus new species 3. In males, the basal plate (Fig. 6-36) and median lobe (Figs. 6-34 and 6-35) are ea sily distinguished from all other species. Description. Male. Form: Elongate oval; contractile; strongly convex dorsally. Length: 1.1 mm (measured from apex of clype us to apex of elytra); breadth: 0.9 mm (measured across elytral humeri). Color: Head, thorax, elytra and underside dark brown, lateral margin of pronotum and posterior margin of elytra translucen t; legs and antennae brown or black. Head: Broad and convex, clypeus produced, narrow, and slightly

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137 reflexed. Eyes large, oblong, with internal margins distinct. Genae not visible from above. Dorsal surface smooth under 100 times magnification, distinctly alutaceous and finely punctate. Antennae 11-segmented including a club with 3 antennomeres, club length about size of the eye. Club flat and distinctly separated from funicle and with a distinctly serrated margin. First cl ub antennomere wider than long, second club antennomere larger than either first or third club antennomere and about as long as wide. Terminal club antennomere truncate (Fig. 6-33 ), setose and about as long as wide. Antennomere 3 as long or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curv ed; anterior angle more narrowly arcuate than posterior. Surface distin ctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1 / 5 Strongly convex, sides s lightly sinuous and apices rounded. Length slightly shor ter than combined width ( 31:38). Uniformly punctate along dorsal surface, smooth at sides and base wi th a large impunctate area at apices of elytra. Appearing alutaceous under 100 times magnification. Median margin and apices of elytra bordered. Underside: Metasternum alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly covering the surface. Legs: Femora glossy, broad, fla ttened and sparsely covered with short hairs. Proand mesofemora about the same width throughout length. All tibiae slightly but distinctly curved and dilated towards ape x. Protibiae with short hairs along outer margin. Mesoand metatibiae with long, stiff hairs along outer margin. Four tarsomeres, claw tarsomere as long or almo st as long as 2 tarsomeres preceding it. Median lobe: Sides parallel or slightly convergent curving into a sharp point (Fig. 6-34).

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138 In profile, strongly curved from base (Fig. 6-35). Median plate on surface very elevated. Basal plate: Sides curving to an evenly rounded apex with a deep, narrow concavity in the center (Fig. 6-36). Female. Unknown. Geographic distribution. Puebla : nr. Tehuacan; Sinaloa : El Dorado (Fig. 6-41). Type material examined. The holotype, deposited in the UCRC, is a male specimen glued to a point with the following labels: MEXICO, Sinaloa, El Dorado, 14-II1929, coll. S. E. Flanders (printed) [white rectangular label]. One male paratype deposited in the UCRC has the same data as the holotype and one male paralectotype EMEC is labelled as follows Mexico: Puebla, 6 km. N Tehuacan, 22-VIII-1987, coll. J. Doyen. Pycnocephalus Sharp 1891 Pycnocephalus Sharp 1891: 373. For a description of the genus, see Smith and Cave (2007). Pycnocephalus metallicus Sharp (Figs. 6-37-6-41) Pycnocephalus metallicus Sharp 1891: 373. Diagnosis. The male is black with a green me tallic sheen on the head, anterior portion of the pronotum, and scape. The female is black and without the metallic sheen. This species is larger (>1.5 mm.) than any of the Cybocephalus in this region. The extremely large and dilated middle and hind tib iae as well as the extremely short, broad head distinguish this species from all other cybocephalids in the Mexico. Description. Male. Form: Elongate, ovate; contractile ; strongly convex dorsally. Length: 1.8-2.2 mm (measured from apex of clypeus to apex of elytra); breadth: 1.3-1.5

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139 mm (measured across elytral humeri). Color: Head and apical portion of pronotum dark but with a green metallic sheen, posterior portion of pronotum, elytra, and underside black, front legs light brown or amber, middl e and hind legs brown or dark brown, scape dark with a blue-green metallic sheen, remaining 10 antennomeres brown or amber. Head: Broad, convex and very short, clypeus extr emely short and broad, with a slightly concave apical margin. Eyes very large, fairly round and oblong with internal margins distinct. Genae not visible from above. Dorsal surface alutaceous and finely punctate. Antenna with 11 antennomeres including a 3-se gmented club about 1/3-1/2 the height of the eye. Club flat, distinctly separated from funicle, and with a distinctly serrated margin. First and second club antennomeres wider than long, terminal club antennomere is the largest club segment and slightly longer than wide, apically rounded and setose (Fig. 637). Third antennomere subequal to antennomeres 4 and 5 combined. Pronotum: Strongly convex, lateral margins curved; ante rior angle more narrowly arcuate than posterior angles. Surface finely punctuate. Scutellum: Triangular with margins slightly convex and sinuous; alutaceous and very sparsely punctured. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex, sides almost parallel and apices rounded, length shorter than combined width (57:70) Distinctly alutaceous and punctate, punctation extending to lateral edges and almost to apices. Median and lateral margins of elytra bordered. Underside: Metasternum alutaceous and roughly punctured, with short coarse hairs uniformly distributed. Abdomina l sternites alutaceous and punctate with long, coarse hairs thinly covering the surface. Legs: Protibiae expanded towards the distal end and setose, profemora narrowing slightly at the distal end. Mesoand metatibiae with long, stiff hairs along the out er margin. Mesoand metafemora and tibiae

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140 very alutaceous, laminiform and greatly dilate d. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres combined. Median lobe: Sides slightly rounded and curving to a gradual rounded tip (F ig. 6-38). In profile, sinuous and curved from middle (Fig. 6-39). Median pl ate on surface slightly elevated. Basal Plate: Sides parallel at base, evenly rounded with 2 small pr otuberances on either side of a flat surface at apex (Fig. 6-40). Female: Similar to male but head, pronotum, and first antennal segment black and without green metallic sheen. Geographic distribution. Mexico: Tamaulipas : Ciudad Victoria Caon La Libertad, Tampico (Fig. 6-41) and Guatemala. Type material examined. The holotype, deposited in the BMNH, is a female specimen glued to a card with Pycnocephalus metallicus, with Cyb being scribbled out and replaced with Pycn, Type D. S. Tama hu, Guat. Champion handwritten on the same card the specimen is glued to. The remaining labels are as follows: TYPE (printed) [white circular label with an orange outer margin] / Tamahu, Vera Paz. Champion (printed) [white rectangular label] / Sp. figure d. (printed) [white rectangular label] / B. C. A. Col. II. l. Pycnocephalus metallicus Sharp / Pycnocephalus metallicus Sharp (handwritten) det. A. Kirejtshuk 1999 (print ed) [white rectangular label] / HOLOTYPE Pycnocephalus metallicus Sharp Det: Trevor Smith (printed ) [red rectangular label]. While this specimen is a syntype, prior to this publication it wa s not a valid holotype, despite the round type label pl aced on the specimen, because this designation was never published. One female paralectot ype is labelled as follows: Guatemala: San Geronimo, Tamahu (BMNH).

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141 Other material examined. Mexico: Tamaulipas, Ciudad Victoria Caon La Libertad, 7-III-1986, coll P. W. Kovarik (1 FSCA); Tampico, 27-?-1912, coll. E. A. Schwarz (1 USNM); Tampico, 26-?-1912, coll. E. A. Schwarz (1, USNM); Tampico, 24-?-1912, coll. E. A. Schwarz (1 USNM); Tampico, 26-?-1912, coll. E. A. Schwarz (1 1 BMNH). Remarks. The type specimen is a female, however enough associated male specimens were available for comparisons a nd a male description including figures of male genitalia to be made.

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142 Figure 6-1. Antenna of C. aciculatus

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143 Figure 6-2. Antenna of Cybocephalus new species 1. Figure 6-3. Median l obe, dorsal view, of Cybocephalus new species 1. Figure 6-4. Median lobe lateral view, of Cybocephalus new species 1. Figure 6-5. Basal plate, ventral view, of Cybocephalus new species 1.

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144 Figure 6-6. Truncate antenna of C. californicus Figure 6-7. Emarginated antenna of C. californicus Figure 6-8. Median l obe, dorsal view, of C. californicus. Figure 6-9. Median lobe lateral view, of C. californicus. Figure 6-10. Basal plate, ventral view, of C. californicus.

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145 Figure 6-11. Collection localities of Cybocephalus aciculatus, Cybocephalus new species 1 and Cybocephalus californicus in Mexico.

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146 Figure 6-12. Antenna of Cybocephalus new species 2. Figure 6-13. Median lo be, dorsal view, of Cybocephalus new species 2. Figure 6-14. Median lobe, lateral view, of Cybocephalus new species 2. Figure 6-15. Basal plate, ventral view, of Cybocephalus new species 2.

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147 Figure 6-16. Antenna of Cybocephalus new species 3. Figure 6-17. Median lo be, dorsal view, of Cybocephalus new species 3. Figure 6-18. Median lobe, lateral view, of Cybocephalus new species 3. Figure 6-19. Basal plate, ventral view, of Cybocephalus new species 3.

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148 Figure 6-20. Antenna of C. nigritulus Figure 6-21. Median lo be, dorsal view, of C. nigritulus. Figure 6-22. Median lobe, lateral view, of C. nigritulus. Figure 6-23. Basal plate, ventral view, of C. nigritulus.

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149 Figure 6-24. Collection localities of Cybocephalus new species 2, Cybocephalus new species 3 and Cybocephalus nigritulus in Mexico.

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150 Figure 6-25. Antenna of C. randalli. Figure 6-26. Median lo be, dorsal view, of C. randalli Figure 6-27. Median lobe, lateral view, of C. randalli Figure 6-28. Basal plate, ventral view, of C. randalli

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151 Figure 6-29. Antenna of C. schwarzi Figure 6-30. Median lo be, dorsal view, of C. schwarzi Figure 6-31. Median lobe, lateral view, of C. schwarzi Figure 6-32. Basal plate, ventral view, of C. schwarzi

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152 Figure 6-33. Antenna of Cybocephalus new species 4. 6-33) antenna; 6-34) median lobe, dorsal view; 6-35) median lobe. lateral view; 6-36) basal plate, ventral view. Figure 6-34. Median lo be, dorsal view, of Cybocephalus new species 4. Figure 6-35. Median lobe, lateral view, of Cybocephalus new species 4. Figure 6-36. Basal plate, ventral view, of Cybocephalus new species 4.

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153 Figure 6-37. Antenna of P. metallicus Figure 6-38. Median lo be, dorsal view, of P. metallicus Figure 6-39. Median lobe, lateral view, of P. metallicus Figure 6-40. Basal plate, ventral view, of P. metallicus

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154 Figure 6-41. Collection localities of Cybocephalus randalli Cybocephalus schwarzi Cybocephalus new species 4 and Pycnocephalus metallicus in Mexico.

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BIOGRAPHICAL SKETCH Trevor Randall Smith was born on February 3, 1977, in Tampa, Florida. The son of an avid outdoorsman, his earliest memories are of hunting, fishing, and hiking in the unique ecosystems of Florida. In December 2000, he graduated from the University of Central Florida with a Bachelor of Science de gree in biology under the guidance of Stuart Fullerton. Trevor was offered an assistantshi p to study entomology at the University of Florida in Summer 2001 and worked with Dr John Capinera. Upon receiving a Master of Science degree in December 2003, Trevor was again offered an assistantship to pursue his Doctor of Philosophy degree at the Universi ty of Florida, this time with advisor Dr. Ronald Cave. 166


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LIFE HISTORY AND PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus
ENDRODY-YOUNGA (COLEOPTERA: CYBOCEPHALIDAE) AND A
TAXONOMIC REVISION OF THE CYBOCEPHALIDAE OF NORTH AMERICA
AND THE WEST INDIES















By

TREVOR RANDALL SMITH


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

UNIVERSITY OF FLORIDA


2006

































Copyright 2006

by

Trevor Randall Smith

































This document is dedicated to Stuart Fullerton without whom none of this would have
been possible. I would also like to dedicate this dissertation to my wife Kathryn who has
been very patient throughout my long academic journey.















ACKNOWLEDGMENTS

First and foremost I would like to offer my gratitude to Ronald D. Cave, my

advisor. He has helped me in more ways than could ever be listed here. I thank Howard

Frank, Michael C. Thomas, Paul E. Skelley, Bill Overholt and Zhenli He for reviews of

this manuscript. A special thanks to Bijan Deghan for offering much needed advice and

discussion on cycads, reviewing my manuscript and serving on my committee even after

his retirement. I also thank Christine Emshousen for tours of and information about the

Montgomery Botanical Center. Simon Yu has given endless help and advice concerning

experimental design of pesticide susceptibility studies, and I also extend my gratitude to

the curators of numerous collections for loans of specimens. Jane Medley and Patricia

Hope have offered tremendous assistance with figures and photographs and I would like

to thank them as well. This research was supported by a grant from the Florida

Department of Agriculture and Consumer Services (DACS 7276186-12).
















TABLE OF CONTENTS



A C K N O W L E D G M EN T S ......... ......... .. ..................................................................... iv

LIST OF TA BLES ............. .. .......................... ........ ...... .. ............. viii

LIST OF FIGURES ......... ......................... ...... ........ ............ ix

A B S T R A C T .......................................... ..................................................x v

CHAPTER

1 LITERATURE REVIEW ..............................................................................1

C y c a d s ........................... ......................................... 1
G general Scale Insect Inform ation ........................................ ........................... 2
Cycad Aulacaspis Scale (CA S) ............................................................................. 2
Chem ical Control of CA S........................................................... .............5
B biological C control of C A S ........................................ ................................. 6
C yb oceph alidae.............................. ............................................. ....................... . 6
Cybocephalids as Biological Control Agents............................................................7
Rhyzobius lophanthae ......... ................................. ......................... ............... 11

2 LIFE HISTORY OF Cybocephalus nipponicus ENDRODY-YOUNGA A
PREDATOR OF Aulacaspis yasumatsui TAKAGI (HOMOPTERA:
D IA SP ID ID A E ) ......... .... ........ .. .. ............ ...................................................... 15

Introduction ................ ........ ................................ ...........................15
M materials and M methods ....................................................................... .................. 16
R results and D discussion ........................................ ................... .. ...... 18

3 PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus AND Rhyzobius
lophanthae (COLEOPTERA: CYBOCEPHALIDAE, COCCINELLIDAE) ............32

In tro d u ctio n ...................................... ................................................ 3 2
M materials and M methods ....................................................................... ..................34
In sects ....................................................................... ................. 3 4
Bioassays Using Coated Glass Vial Method.....................................................35
Statistical A naly sis ......................... .. .................... ......... ........... 36
R e su lts ...................................... .................................................... 3 7









Effects of Pesticide on C. nipponicus.....................................................37
Effects of Pesticide on R. lophanthae ....................... ........ ............ 37
D isc u ssio n ............................................................................................................. 3 7

4 THE CYBOCEPHALIDAE (COLEOPTERA) OF AMERICA NORTH OF
M E X IC O .......................................................4 2

In tro d u ctio n .......................................................................................4 2
Taxonom ic History ......................... ........................ ..... ..... 45
M materials and M methods ....................................................................... ..................46
M a te ria ls ...................................................................................................4 6
M e th o d s ...........................................................................4 7
D definitions ..........................................................................48
Cybocephalus Erichson 1844 ............................................................49
Key to the species of Cybocephalus of America north of Mexico ...........................49
Cybocephalus californicus Horn ......................................................... 50
Cybocephalus kathrynae T. R. Smith, New Species..........................................60
Cybocephalus nigritulus LeConte ................. ................. .............................63
Cybocephalus nipponicus Endrdy-Younga ..................................... ...............68
Cybocephalus randalli T. R. Smith, New Species .......................................... 73

5 THE CYBOCEPHALIDAE (COLEOPTERA) OF THE WEST INDIES AND
T R IN ID A D ........................................................................................................... 9 1

In tro d u ctio n ................................................................... ................................ 9 1
M materials an d M eth od s ......................................................................................... 92
M materials ...................................... ................................................ 9 2
M e th o d s ...........................................................................9 3
Definitions ........................................ ........................93
Key to the Cybocephalidae of the West Indies and Trinidad .............. ............... 93
Cybocephalus Erichson 1844 ........................................................... ................. 94
Cybocephalus antilleus T. R. Smith, New Species ............................................ 94
Cybocephalus caribaeus T. R. Smith, New Species ..................................... 96
Cybocephalus iviei T. R. Smith, New Species ........................................ ....99
Cybocephalus nipponicus Endr6dy-Younga .................................... .....102
Cybocephalus geoffreysmithi T. R. Smith, New Species ......................................103
Pycnocephalus Sharp 1891................................ ........................105
Pycnocephalus deyrollei (Reitter), New Combination ............................................106

6 THE CYBOCEPHALIDAE (COLEOPTERA) OF MEXICO .............................119

In tro d u c tio n ......................................................................................................... 1 1 9
M material and M methods .......................................................................... ............... 119
M ateria ls .......................................................................................... 1 19
M e th o d s ................... .............................................................................. 12 0
D definitions ............................................................0... ......120
Key to the Cybocephalidae of Mexico ...................................................121









Cybocephalus Erichson 1844 .................................................. ..... ... ........... 122
Cybocephalus aciculatus Champion..................................... ........................ 122
Cybocephalus N ew Species 1 ........................................... ............................ 124
Cybocephalus californicus H orn ........................................ ......................... 126
Cybocephalus N ew Species 2................................. ............... ............... 128
Cybocephalus New Species 3 ....... ................... ............... 129
Cybocephalus nigritulus LeConte ........................................ ........................ 131
Cybocephalus randalli T. R. Smith ........................ ............................ 132
Cybocephalus schwarzi Champion...................................... ......................... 134
Cybocephalus N ew Species 4................................. ............... ............... 136
P y cnocep halus Sharp 189 1......................................... ........................................ 138
Pycnocephalus m etallicus Sharp ........................................ ......................... 138

L IST O F R E FE R EN C E S ..................... .. .. ......... .. .......................... ........................155

BIOGRAPHICAL SKETCH .............. ............................................................. 166
















LIST OF TABLES


Tablege

2-1 Development of Cybocephalus nipponicus on Aulacaspis yasumatsui .........3.....1..

2-2 Mortality (%) of immature stages of Cybocephalus nipponicus during rearing on
Aulacaspisyasumatsui....................................... 31

3-1 Field rates (IX) for each pesticide used. ....................................... ............... 40

3-2 Percent mortality of Cybocephalus nipponicus per 30 individuals exposed. X =
fie ld ra te ...................................... .................................. ................ 4 0

3-3 Percent mortality ofRhyzobius lophanthae per 30 individuals exposed. X = field
rate..................... .......................................40

3-4 Student-Newman-Keuls test showing ranked values of mortality of adult
Cybocephalus nipponicus and Rhyzobius lophanthae. .........................................41
















LIST OF FIGURES


Figurege

2-1 Dorsal habitus. A, Cybocephalus binotatus. B, Cybocephalus nipponicus. ...........25

2-2 Egg of Cybocephalus nipponicus. ........................................ ........................ 25

2-3 First instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral
h ab itu s. ........................................................... ................ 2 6

2-4 Short trumpet-shaped setae on third instar larva of Cybocephalus nipponicus. ......26

2-5 Third instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral
oblique habits. .......................................................................27

2-6 Pupal chambers of Cybocephalus nipponicus. A, Pupal chamber made from
female scale covers. B, Pupal chamber made from sand. ......................................28

2-7 Pupa of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral habitus ............29

2-8 Adult C. nipponicus, lateral habitus ....................................................................... 30

2-9 L arval m ortality over tim e........................................................................... .. ..... 30

4-1 Complete Cybocephalus nipponicus genitalia: ED = ejaculatory duct; IS =
internal sac; MA = muscle attachment; ML = median lobe; MS = median strut;
Tbp = tegmen basal plate; Tdp = tegmen dorsal piece; TS = tegminal strut. ..........77

4-2 Lateral habitus of Cybocephalus randalli. .................................... .................77

4-3 Truncate antenna of C. californicus. ............................................. ............... 78

4-4 Emarginate antenna of C. californicus .......................................... ............... 78

4-5 Median lobe, dorsal view, of C. californicus.. ................... ........................ 78

4-6 Median lobe, dorsal view (slide mounted), of C. californicus ..............................78

4-7 Median lobe, lateral view, of C. californicus. ................... ......................... 78

4-8 Basal plate, ventral view, of C. californicus. ....................................... ............... 78









4- 9 U.S. states and Canadian provinces from which specimens of Cybocephalus
californicus have been collected. ........................................ ........................ 79

4-10 Scutellum of Cybocephaus nipponicus. ...................................... ............... 80

4-11 Scutellum of Cybocephalus kathrynae................................. ............... 80

4-12 A antenna of C. kathrynae. ........................................ ................................. 81

4-13 Median lobe, dorsal view (same as slide mounted), of C. kathrynae .......................81

4-14 Median lobe, lateral view, of C. kathrynae. ................................... ...............81

4-15 Basal plate, ventral view, of C. kathrynae. ................................... ............... 81

4-16 Collection localities of Cybocephalus kathrynae in Florida .............. ...............82

4-17 A antenna of C. nigritulus ....................................................................... 83

4-18 Median lobe, dorsal view, of C. nigritulus..........................................................83

4-19 Median lobe, dorsal view (slide mounted), of C. nigritulus. .............. ...............83

4-20 M edian lobe, lateral view, of C. nigritulus. .................................. .................83

4-21 B asal plate, ventral view of C. nigritulus ........................................ .....................83

4-22 U.S. states and Canadian provinces from which specimens of Cybocephalus
nigritulus have been collected ..................................................................... ..... 84

4-23 A ntenna of C nipponicus ........................................................................... ... .... 85

4-24 Median lobe, dorsal view (same as slide mounted), of C. nipponicus ....................85

4-25 M edian lobe, lateral view, of C. nipponicus. ................................... ..................... 85

4-26 Basal plate, ventral view, of C. nipponicus......................................................85

4-27 U.S. states from which specimens of Cybocephalus nipponicus have been
collected. ............................................................................86

4-28 D orsal habits of C binotatus .......................... .. ................................................ 87

4-29 Dorsal habitus of C. nipponicus. ........................................ ......................... 87

4-30 A antenna of C randalli. .................................................................... ..................88

4-31 Median lobe, dorsal view, of C. randalli. ..................................... ............... 88









4-32 Median lobe, dorsal view (slide mounted), of C. randalli ......................................88

4-33 Median lobe, lateral view, of C. randalli. ..................................... ............... 88

4-34 Basal plate, ventral view, of C. randalli. ..................................... ............... 88

4-35 Striations at the apices of elytra on C. randalli .................. ...............89

4-36 U.S. states from which specimens of Cybocephalus randalli have been
collected. ............................................................................90

5-1 V central habitus of C. nipponicus ................................................ ....... ........ 110

5-2 Ventral habits of P. deyrollei. ....................................................... ........... 110

5-3 A antenna of C antilleus................................................................ ..................... 1

5-4 M edian lobe, dorsal view, of C. antilleus. .......... .........................................111

5-5 Median lobe, lateral view, of C. antilleus. ........ ..... ........... ....................111

5-6 Basal plate, ventral view, of C. antilleus ............... ....................... ............111

5-7 A antenna of C caribaeus.............................................................. ............... 112

5-8 Median lobe, dorsal view, of C. caribaeus. ............ ................................... 112

5-9 Median lobe, lateral view, of C. caribaeus. ............................................. 112

5-10 Basal plate, ventral view, of C. caribaeus............................ .. ............... 112

5-11 A antenna of C. iviei. ................................................... .... .. ........ .. .. 113

5-12 Median lobe, dorsal view, of C. iviei. ......................................... ...............113

5-13 M edian lobe, lateral view, of C. iviei. ........................................... ............... 113

5-14 Basal plate, ventral view, of C. iviei. ............................................ ............... 113

5-15 Scutellum of C. iviei .................. .......................... .. ...... ................ 114

5-16 Scutellum of C nipponicus. ....................................................................... ....... 114

5-17 Invaginations at the elytral apices of C. iviei. .......................................................115

5-18 A ntenna of C. nipponicus ........................................................................ 116

5-19 M edian lobe, dorsal view of C. nipponicus.........................................................116

5-20 M edian lobe, lateral view, of C. nipponicus. ....................................................... 116









5-21 Basal plate, ventral view of C. nipponicus.......................................................... 116

5-22 A ntenna of C. geoffreysm ithi. .................................................................... ......117

5-23 M edian lobe, dorsal view, of C. geoffreysmithi. .................................................... 117

5-24 Median lobe, lateral view, of C. geoffreysmithi ............................... ............... 117

5-25 Basal plate, ventral view, of C. geoffreysmithi. ..................................................... 117

5-26 Antenna of P. deyrollei. ............. ............... .............................................118

5-27 Hind leg of P. deyrollei. .................................................... ...............118

5-28 M edian lobe, dorsal view, of P. deyrollei. ............. ............................... ......... 118

5-29 Median lobe, lateral view, of P. deyrollei........................ ...................118

5-30 Basal plate, ventral view, of P. deyrollei. ........................................ ...............118

6-1 Antenna of C. aciculatus....................... ................................... 142

6-2 Antenna of Cybocephalus new species 1. .................................... .....................143

6-3 Median lobe, dorsal view, of Cybocephalus new species 1.............................. 143

6-4 Median lobe, lateral view, of Cybocephalus new species ................................. 143

6-5 Basal plate, ventral view, of Cybocephalus new species 1. .................................. 143

6-6 Truncate antenna of C. californicus. ........................................... ............... 144

6-7 Emarginated antenna of C. californicus. ................................... ............... 144

6-8 Median lobe, dorsal view, of C. californicus. ................................................ 144

6-9 M edian lobe, lateral view, of C. californicus.................................. ...............1. 44

6-10 Basal plate, ventral view, of C. californicus. ................................................. 144

6-11 Collection localities of Cybocephalus aciculatus, Cybocephalus new species 1
and Cybocephalus californicus in Mexico. ...................................................145

6-12 Antenna of Cybocephalus new species 2. .................................... .....................146

6-13 Median lobe, dorsal view, of Cybocephalus new species 2............................. 146

6-14 Median lobe, lateral view, of Cybocephalus new species 2.............................. 146

6-15 Basal plate, ventral view, of Cybocephalus new species 2 .................................. 146









6-16 Antenna of Cybocephalus new species 3. ...... ........... .. ......... ..................147

6-17 Median lobe, dorsal view, of Cybocephalus new species 3.............................. 147

6-18 Median lobe, lateral view, of Cybocephalus new species 3.............. ...............147

6-19 Basal plate, ventral view, of Cybocephalus new species 3 ............................... 147

6-20 A ntenna of C nigritulus ........................................................................... ....... 148

6-21 M edian lobe, dorsal view, of C. nigritulus.............. ..........................................148

6-22 Median lobe, lateral view, of C. nigritulus. ................................................ 148

6-23 Basal plate, ventral view, of C. nigritulus.............. ............................................148

6-24 Collection localities of Cybocephalus new species 2, Cybocephalus new species
3 and Cybocephalus nigritulus in M exico.................................. ............... 149

6-25 A antenna ofC randalli. ........................................... ......................................... 150

6-26 Median lobe, dorsal view, of C. randalli. ................................................... 150

6-27 Median lobe, lateral view, of C. randalli. ................................... ............... 150

6-28 Basal plate, ventral view, of C. randalli. ............. ........................ ......... ...... 150

6-29 A antenna of C. schwarzi. ............................................... .............................. 151

6-30 Median lobe, dorsal view, of C. schwarzi.........................................................151

6-31 M edian lobe, lateral view, of C. schwarzi ............... ......... .. ............... ....... 151

6-32 Basal plate, ventral view, of C. schwarzi. ................................... .....................151

6-33 Antenna of Cybocephalus new species 4. 6-33) antenna; 6-34) median lobe,
dorsal view; 6-35) median lobe. lateral view; 6-36) basal plate, ventral view.......152

6-34 Median lobe, dorsal view, of Cybocephalus new species 4............................. 152

6-35 Median lobe, lateral view, of Cybocephalus new species 4............................. 152

6-36 Basal plate, ventral view, of Cybocephalus new species 4 .................................. 152

6-37 A ntenna of P. m etallicus. .............................................. ............................. 153

6-38 Median lobe, dorsal view, of P. metallicus. .................... ......................... 153

6-39 Median lobe, lateral view, of P. metallicus.......... ................................ 153









6-40 Basal plate, ventral view, of P. metallicus. ....................................................... 153

6-41 Collection localities of Cybocephalus randalli, Cybocephalus schwarzi,
Cybocephalus new species 4 and Pycnocephalus metallicus in Mexico. ..............154















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

LIFE HISTORY AND PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus
ENDRODY-YOUNGA (COLEOPTERA: CYBOCEPHALIDAE) AND A
TAXONOMIC REVISION OF THE CYBOCEPHALIDAE OF NORTH AMERICA
AND THE WEST INDIES

By

Trevor Randall Smith

December 2006

Chair: Ronald D. Cave
Major Department: Entomology and Nematology

The life history of the predatory beetle Cybocephalus nipponicus Endr6dy-

Younga was studied by rearing the beetle on the cycad aulacaspis scale, Aulacaspis

yasumatsui Takagi. Mean developmental times of egg (7.3 0.8 days), larval (13.7 + 1.1

days), and pupal (18.6 + 1.6 days) stages were determined. Mortality in each life stage,

adult longevity, and adult sex ratios also were measured. A clarification of differences

between C. nipponicus and C. binotatus Grouvelle is included.

The susceptibility of the predatory beetles C. nipponicus Endr6dy-Younga and

Rhyzobius lophanthae Blaisdell to six pesticides commonly used for treating cycad

aulacaspis scale was tested. Three concentrations (half field rate, field rate, and twice

field rate) of each pesticide were tested against both beetle species using a coated glass

vial bioassay. Nearly 100% mortality in both beetle species occurred at all

concentrations when treated with methidathion, dimethoate, and malathion. Insecticidal









soap, fish oils, and imidacloprid were much less toxic. At half the simulated field rate, C.

nipponicus had 33% survivorship with insecticidal soap, 23% survivorship with

imidacloprid, and 17% survivorship with fish oil. At half the simulated field rate, R.

lophanthae had 56% survivorship with insecticidal soap, 36% survivorship with

imidacloprid, and 53% survivorship with fish oil. Mortality rate for each beetle species

rose with increasing concentration of each pesticide.

The 17 species of Cybocephalidae in North America (including Mexico) and the

West Indies are revised. Included are redescriptions of Cybocephalus aciculatus

Champion, C. californicus Horn, C. nigritulus LeConte, C. nipponicus Endrody-Younga,

C. schwarzi Champion, Pycnocephalus metallicus Sharp and a new combination of P.

deyrolli (Reitter). Also included are descriptions of 10 new species: C. antilleus, C.

caribaeus, C. iviei, C. kathrynae, C. randalli, C. geoffreysmithi and four as yet unnamed

new species from Mexico. A key to species, illustrations of morphological features

including detailed drawings of male genitalia, distribution data, and host lists are

provided. The confusion involving C. nipponicus and C. binotatus Grouvelle is

discussed and the differences between them are made evident.














CHAPTER 1
LITERATURE REVIEW

Cycads

Cycads are an ancient group of plants, sometimes called the coelacanthss of the

plant world," that date back to the Paleozoic era (Moretti 1990). However, the true rise

and dominance of the cycads occurred during the Mesozoic era. Cycads are believed to

be an evolutionary intermediate between ferns and flowering plants (Whiting 1962).

Cycads belong to three families: Cycadaceae which includes a single genus, Cycas;

Stangeriaceae which also contains only 1 genus, Stangeria; and Zamiaceae which

contains 8 genera, Bowenia, Ceratozamia, Dioon, Encephalartos, Lepidozamia,

Macrozamia, Microcycas, and Zamia (Whitelock 2002). There are almost 300 species of

cycads worldwide, most of which are found in tropical and subtropical environments

(Hill 2004). Many of these species are endangered or threatened, with this threat due

more to determined collectors than to deforestation, agriculture, and urban sprawl (Giddy

1990).

While in some cases cycads are used for food or fertilizer, their chief economic

importance is as ornamental landscape plants (Thieret 1958). These plants can be found

in hundreds of nurseries all over the state of Florida. While older and consequently larger

plants can be quite expensive, smaller 1 gallon pot size C. revoluta can sell for as little as

$10.00 (Home Depot). Aside from its natural beauty and hardiness, Cycas revoluta is a

very popular plant with growers because it can be propagated through cutting lateral

outgrowths or "pups".









General Scale Insect Information

The armored scale insects (Diaspididae) belong to the superfamily Coccoidea in the

order Hemiptera. There are over 1,800 described species of armored scale worldwide

(Ben-Dov 1993) and at least 300 occur in North America (Borror et al. 1989). Many of

these scales are pests of agriculture and horticulture, generally weakening plants through

sap feeding and often causing excessive sooty mold growth on excreted honeydew. In

their native habitats scale insects are usually controlled by natural predators and

parasitoids (Hodgson and Martin 2001). The soft-bodied females and early-instar males

of the tribe Diaspidini live beneath a scale covering made of wax secreted by the insect

and mixed with shed exuviae of earlier instars. The scale covering of males is usually

smaller and more elongate than that of the females. The first instar, or "crawler," stage is

mobile and able to spread to other plants. Female crawlers will settle and insert their

mouthparts into the plant, becoming sessile and remaining in that state throughout their

lifetime. Females will molt twice after the crawler phase, becoming an adult in the third

instar. Males become sessile after the crawler phase and remain in that state through the

second and third instar before molting into a fourth prepupa stage. The emergent fifth

instar male is winged and without mouthparts (Hamon 2000).

Cycad Aulacaspis Scale (CAS)

At least 20 species of scale insect occur on cycads in Florida, 19 of which cause

very little damage to cycads (Dekle 1976). The cycad aulacaspis scale (CAS),

Aulacaspis yasumatsui Takagi, is the most damaging scale found on cycads in Florida

(Hodges et al. 2003). This scale is native to Southeast Asia. It has also been found on

several islands in the Caribbean, as well as Hawaii (Ben-Dov et al. 2003). In 1992 A.

yasumatsui became such a problem in Hong Kong that 70-100% mortality of Cycas









revoluta Thunberg was recorded (Hodgson and Martin 2001). In Guam the endemic

Cycas micronesica K.D. Hill has been severely affected by this damaging scale (R.

Muniappan 2005, personal communication). The first detection of the CAS in Florida

occurred in 1996 in Miami at the Montgomery Botanical Center. The scale is thought to

have come in on infested cycads imported from Southeast Asia (Howard and Weissling

1999). By the end of 1997 the scale had spread throughout Miami and as far north as

Lake Okeechobee and could be found on 20 species of cycads (Howard and Weissling

1999); however it seemed to prefer the genera Cycas and Stangeria, a very rare genus.

This led to the spread of the scale to Hawaii in 1998 through the legal importation of

infested cycads from Florida (Hodgson and Martin 2001). Currently CAS has been

reported from Pensacola east to Jacksonville and south into the Florida Keys. How far

north the scale is actually established is not known. It is suspected that many of the

infested cycads in north Florida were in fact transplants from southern nurseries rather

than the pest's natural progression north. However, this is just speculation.

Aulacaspis yasumatsui was first described by Dr. Sadao Takagi from specimens

collected in Bangkok, Thailand (Takagi, 1977). This scale will feed on a large number of

cycads; however, the most commercially significant are those of the genus Cycas. In

Florida, C. revoluta, the king sago, and Cycas rumphii Miq., the queen sago, are the most

popular cycads used in landscaping, and unfortunately, both of these cycads are severely

attacked by the cycad scale (Howard et al. 1999). Mature A. yasumatsui females have a

white armor 1.2-1.6 mm in diameter that is usually of a pyriform shape common in the

tribe Diaspidini, with the exuviae at one end (Ben-Dov 1990). This is by no means the

only shape in which this scale may appear. The females of this species may have a large









number of shapes and sizes. The male has a very typical, of the tribe Diaspidini,

tricarinate, 0.5-0.6 mm-long, white teste with the exuviae at the cephalic end (Howard et

al. 1999).

In the field the magnolia white scale, Pseudaulacaspis cockerelli (Cooley), often

found on cycads, is frequently confused with CAS. However, once the covers are

removed major differences in the two species can be observed. All stages of the CAS life

cycle, from egg to adult, are a uniform orange color, whereas all stages of the magnolia

white scale are yellow. Also, the female CAS has a swollen prosoma and is quite

compact, whereas the magnolia white scale has a slender prosoma and is relatively

elongate (Hodges et al. 2003). The magnolia white scale populates more heavily on the

adaxial surface, while CAS creates extremely dense populations on the abaxial surface

with relatively few individuals settling on the upper surface (Howard and Weissling

1999). Most importantly, when a cycad is heavily infested with CAS, the sheer volume

of individual scales can become so great that the entire plant is coated with a white crust

usually made up mostly of male scales (Howard et al. 1996). By contrast, magnolia

white scale infestations are much less severe (Howard and Weissling 1999). Lastly, CAS

infests all parts of the cycad including the leaves, cones, fruits, megasporophylls, stems

and roots (Howard and Weissling 1999), whereas the magnolia white scale attacks only

the leaves.

Before the introduction of CAS the only scale of the genus Aulacaspis in Florida

was Aulacaspis rosae (Bouche), a non-native pest of roses (Dekle 1976). Aulacaspis

rosae may be a close relative of A. yasumatsui because the 2nd instar of A. rosae is very









similar morphologically to that ofA. yasumatsui (Takagi 1977), but with such different

host plants these two scales are not often confused.

Chemical Control of CAS

There has been some success controlling the cycad aulacaspis scale with various

pesticides. Oils, either an ultra-fine horticultural oil or a product containing fish oil, seem

to be the most effective chemical control method (Hodges et al. 2003). This is not really

surprising given that oils have long been used to control armored scale insects. The oil

not only covers the insects and suffocates them but also covers the surface of the plant

making it difficult for crawlers to settle onto the plant (Howard and Weissling 1999).

The proper application of the oils is difficult due to the scale's tendency to heavily infest

the abaxial surface of the leaves, which is difficult to spray (Howard and Weissling

1999). In the case of C. revoluta, the architecture of the plant itself, with the margins of

the leaflets curling down and inward and forming a trough on the abaxial surface of the

leaflet, makes foliar oil treatments difficult (Hodges et al. 2003). Frequent (every two

weeks) or "as needed" use of oils seems to be the most effective technique for controlling

this scale, and by mixing treatments of oil with treatments of contact insecticides such as

malathion or carbaryl, even greater scale mortality can be achieved (Hodges et al. 2003).

Horticultural oils also seem to at least help control CAS on the root systems of potted

cycads. Hodges et al. (2003) found that drenching the roots of an infested cycad in 2%

horticultural oil resulted in 100% mortality of mature females on the roots. However, a

root drench would be very difficult to accomplish properly on field-grown cycads. The

use of systemics such as methidathion and dimethoate has yielded mixed results, being

very effective in some cases and completely ineffective at controlling the scale in other

cases (Hodges et al. 2003). Imidacloprid as a soil drench can be very effective but









Howard and Weissling (1999) found that this product had to be mixed at such high

concentrations that not only was the product not labeled for such high rates but also the

whole process became extremely uneconomical. They were completely unable to control

CAS using imidacloprid at label rates. Insect growth regulators such as pyroproxyfen,

sold under the trade name Distance, have met with some success controlling CAS

(Emshousen and Mannion 2004). Homeowners have found that liberal application of

soap can be a very effective control method as well (personal observation, 2004).

Biological Control of CAS

The cycad aulacaspis scale is considered a pest in Thailand, but native parasitoids

are reported to control its populations (Tang et al. 1997). Howard et al. (1999) reported

the lack of any native natural enemies in Florida as one of the major reason for the rapid

spread of CAS. For this reason two natural enemies were imported from Thailand and

released by Dr. Richard Baranowski of the UF/IFAS Tropical Research and Education

Center at Homestead in 1998. These natural enemies were a parasitic wasp, Coccobius

fulvus (Compere and Annecke), and a predacious beetle, which at the time was identified

as Cybocephalus binotatus Grouvelle. In Hawaii the coccinellid Rhyzobius lophanthae

(Blaisdell) was found to feed quite readily on CAS (Heu and Chun 2000). Howard

(1997) mentions the use of R. lophanthae in Florida as a potential predator of CAS.

Cybocephalidae

The Cybocephalidae differ greatly from the Nitidulidae, the family in which they

have been historically placed, not only because they are predatory but also because their

basic morphology and anatomy are quite different. Cybocephalid adults have a 4-4-4

tarsal formula instead of 5-5-5 found in Nitidulidae. There are 5 visible ventral plates

(leaving out the male anal plate) and 5 abdominal spiracles in cybocephalids instead of









the 6 and 6 that occur in nititdulids. The body of cybocephalids is retractile allowing the

mandibles in repose to rest against the metasternum, unlike any other nitidulid. The

larvae of Cybocephalidae have a head without dorsal sutures, lack pregomphi and

urogomphi on abdominal tergite XI, and have hypostomal rods with divergent

hypostomal ridges present posteriorly, hypopharynx without a sclerome and bracons,

maxillae without mola, and annular spiracles with 2 lateral air tubes. In contrast, the

larvae of Nitidulidae have pregomphi and urogomphi, no hypostomal rods but with

hypostomal ridges strongly convergent posteriorly, hypopharynx with a sclerome and

bracons, maxillae with raised mola, and biforous spiracles (Kirejtchuk 1997). This has

led to the cybocephalids being shuffled around between family (Murray, 1864, Parsons

1943, Endr6dy-Younga 1968) and subfamily (Horn 1879, Arnett 1960, Habeck 2002)

status.

Before the taxonomic study presented here, only one species of Cybocephalidae,

Cybocephalus nigritulus LeConte, was known to be native to the state of Florida; a

second species is described in Chapter 4. Howard and Weissling (1999) reported C.

binotatus was released and has subsequently become established in south Florida.

However, as will be shown later, this species was misidentified and is actually another

Asian species, Cybocephalus nipponicus Endr6dy-Younga. Cybocephalus nipponicus

has been widely used in the United States as a biological control agent for the euonymus

scale, Unaspis euonymi (Comstock), and for many other pest scale species around the

world.

Cybocephalids as Biological Control Agents

With the exception of coccinellids, species of the genus Cybocephalus are the most

important predators of armored scales (Blumberg and Swirski 1982). While









cybocephalids are well-known armored scale feeders, their effectiveness as biological

control agents have not been well studied (Drea and Carlson 1988). Even with such a

limited understanding of the biology and ecology of this group of beetles, species of

Cybocephalus are being released in many areas of the world. There is very little

literature on cybocephalids and what there is usually consists of a small note about them

being released and no evaluation data (Labuschange et al. 1996; Swirski and Wysoki

1995; Hodges et al. 2003). In South Africa, C. nipponicus, along with a parasitoid

Aphytis sp., was imported from Thailand and successfully established in commercial

mango orchards. In the following four seasons these beneficial insects seemed to

effectively control the mango scale, Aulacaspis tubercularis Newstead, with

augmentative releases. A similar release of C. binotatus (but most likely C. nipponicus)

to control the Japanese bayberry whitefly, Parabemisia myricae (Kuwana), on citrus and

avocado trees in Israel was less successful. In this case the researchers were unable to

establish a viable population of the predatory beetle in the field (Swirski and Wysoki

1995).

Drea and Carlson (1988) and Van Driesche et al. (1998) were able to establish C.

nipponicus in the Virginia/Maryland area as well as in New England. In 1995, the New

Jersey Department of Agriculture embarked on an aggressive plan to combat the

euonymus scale using C. nipponicus as a biological control agent (Hudson et al. 2001).

By the year 2000, after many supplemental release and recovery plans had been

implemented, it was obvious that the beetles had established populations all over the state

where they were controlling and in some cases eradicating the euonymus scale.









Others have noted cybocephalids acting as a natural control of diverse species of

armored scales, including at least eight species in Turkey (Erler and Tung 2001), coconut

scale (Aspidiotus destructor (Signoret)) in Brazil (Lima 2002), and the brown apricot

scale (Lecanium corni Bouche) and San Jose scale (Quadraspidiotusperniciosus

(Comstock)) (Heintz 2001) in California. However, in Mauritius the native Cybocephalus

mollis Endr6dy-Younga seemed unable to control the spread of the sugarcane scale,

Aulacaspis tegalensis (Zhnt.), even when in conjunction with three other natural enemies

of the scale (Williams and Greathead 1973).

Because many scale insects persist year-round, it is important that a biological

control agent be found that is also persistent throughout the year. The cybocephalids are

uniquely suited for this in that the placement of eggs and subsequent development of

larvae beneath the armored scale allow them some protection from both the elements and

pesticides (Alvarez and Van Driesche 1998a). In Greece, Katsoyannos (1984) found that

Cybocephalusfodori Endrody-Younga was able to survive in pesticide-treated fruit

orchards. In date palm plantations in Israel, Kehat et al. (1974) found that while all

coccinellids in a chemically treated plantation died, species of Cybocephalus survived.

The dietary needs of cybocephalids also make them good candidates for use as

biological control agents. Alvarez and Van Driesche (1998a) found that at low scale

densities cybocephalids were able to maintain their populations and keep euonymus and

San Jose scale populations in check. In the case of C. nipponicus, an average of 19.5

scales were attacked over the entire larval lifetime, contrasted with an average of 199

green scales (Ceroplastesjaponicus Green) consumed by the coccinellid Chilocorus

kuwanae Silvestri (Xia et al. 1986). Thickness of the scale cover is also a factor in









effectiveness of a beetle as a predator. Blumberg and Swirski (1982) found that C.

micans and C. nigreceps were unable to feed on adult female diaspidid scales. It has also

been noted that coccinellids are more likely to feed on scales with thin or easily

penetrated scale covers (Honda and Luck 1995). Cybocephalus nipponicus seemed to be

less affected by scale cover characteristics (Alvarez and Van Driesche 1998a). In fact, C.

nipponicus was able to feed on the adult females of both the San Jose scale and the

euonymus scale (Alvarez and Van Drieche 1998a). In the absence of prey, female

cybocephalids are able to withhold eggs for up to 2 days, indicating that oviposition

strategy is not only governed by food consumption but by some qualitative features of the

scale population (Alvarez and Van Drieche 1998b). In the presence of greater scale

densities cybocephalids will increase their egg production until they reach a constant.

Again this allows cybocephalids to maintain their populations at low scale densities. If

circumstances allow, female cybocephalids will lay only one egg under a single scale

cover. However, if the number of scales in a patch is low and the beetle cannot find new

scales it will lay eggs under a scale that has already been parasitized (Alvarez and Van

Drieche 1998b). The beetle larvae consume more prey when feeding on young scales;

therefore it seems that for greater survivability of their offspring female cybocephalids

prefer to oviposit on older scales (Alvarez and Van Driesche 1998b). Alvarez and Van

Driesche (1998b) found that the highest larval survival rate could be found in larvae

feeding on scales older than 30 days. It was also noted by Blumberg and Swirski (1982)

that C. nigriceps very rarely laid eggs under dead female scales.

Certain aspects of the biology of Cybocephalus nigriceps (J. Sahlberg),

Cybocephalus micans Reitter, and Cybocephalus gibbulus Erichson were studied by









Nohara and Iwata (1988) and Blumberg and Swirski (1982). Tanaka and Inoue (1983)

were the first to really study the feeding behavior of C. nipponicus. Later, Drea and

Carlson (1988) and Alvarez and Van Driesche (1998a, 1998b) continued the biology of

C. nipponicus. There remains much to be studied about the life cycle, breeding habits,

oviposition behavior, or dietary requirements of C. nipponicus to determine whether it

has any significant effect as a biological control agent of the cycad aulacaspis scale.

Without some form of baseline data on the beetle, no real effectiveness research can be

done.

Rhyzobius lophanthae

Rhyzobius (= Lindorus) lophanthae (Blaisdell) is a small, pubescent coccinellid

belonging to the tribe Coccidulini. The adult is 2.4-2.5 mm in length and between 1.7

and 1.8 mm in width, and has black or brown elytra and an orange-brown thorax and

head region. The body form is elongate or oval with a dense mat of hairs covering the

dorsal surface. The head is partly concealed beneath the pronotum, with 11-segmented

antennae, the last 3 segments of which are broader then the rest to form a club. The tarsal

claws are not toothed. This beetle is often referred to as the singular black lady beetle or

the scale destroyer.

Rhyzobius lophanthae is a coccoidophagous predator native to Australia. It is

considered by many to be one of the most economically important natural enemies of

armored scale insects (Yus 1973, Rosen 1990, Stathas 2001). This beetle has been

released in many areas of the world to control a plethora of armored scale species (Honda

and Luck 1995). There are many examples of successful control of scale insects using R.

lophanthae, especially in the Mediterranean region (Greathead 1973). There have been

some very high profile failures as well, such as the inability ofR. lophanthae to control









the California red scale in California (Greathead 1973). Honda and Luck (1995) found

that the morphological characteristics of the scale itself are a major determining factor in

how effective R. lophanthae will be in controlling a scale species. They discovered that

Rhyzobius mandibles were not as effective as those of other species of coccinellids that

specialize on armored scales, such as Chilocorus cacti (L.), a species frequently seen on

scale-infested cycads in Florida (RD Cave 2006, personal communication). Rhyzobius

lophanthae lacks a tooth at the apex used for prying scales from the substrate and the

incisor region is neither as acutely angled nor as sharp as that of specialists like C. cacti.

Being much less specialized and more of a generalist predator has allowed R.

lophanthae to be used to control a large number of pest species. Rhyzobius lophanthae

seem to be especially effective in closed environments such as greenhouses or in

botanical gardens (Anonymous 2003). They have been recorded feeding on many insects

other than scales including aphids, small caterpillars, whitefly, mites, thrips, psyllids,

mealybugs, and other soft bodied insects and their eggs. These beetles are also voracious

feeders of the immature stages and eggs of scale insects.

The first introduction ofR. lophanthae into the United States was initiated by

Albert Koeble and releases occurred between 1889 and 1892 (Greathead 1973). It failed

to control Saissetia oleae (Olivier), but did establish itself as a predator of armored scales

on citrus and eventually spread throughout the rest of the United States. It was

introduced into Hawaii from California in 1894 (Heu et al. 2003) and has been recorded

feeding on CAS (Hara et al. 2004, personal observation). The larvae and adults have

been observed feeding on eggs, immature, and adult CAS. This beetle was imported and

officially released in Florida to control oleander scale, Aspidiotus nerii Bouche, and other









"scale insects" after 1982 (Frank and McCoy 1994). However, many specimens in the

Florida State Collection of Arthropods (FSCA) were collected in the state as early as

1936, indicating that this beetle has been established in Florida more than 70 years.

Interestingly, on scale-infested cycads in Florida it has been seen only in downtown

Tampa and on the campus of Florida State University in Tallahassee.

Following emergence from the egg, the small, gray larvae immediately begin to

feed on scale eggs and crawlers, which are much more numerous than adult scales, but

will feed on the adult scales as well. The larvae pass through 4 instars, growing to

around 3 mm before pupation. The beetles will often pupate on the plant where the food

source is found. At 25C, the total development of the life cycle ofR. lophanthae takes

about 34 days (Stathas et al. 2002). The adult beetles can live up to 9 months but average

between 5 and 6 (Stathas 2000).

These beetles are exceptional biological control agents because of their high

fecundity, lack of parasitoids, the absence of diapause, and their resistance to low

temperature especially in the immature stages (Rubstov 1952, Smimoff 1950, Stathas

2000). Female R. lophanthae are able to lay hundreds of eggs in a lifetime. Stathas

(2001) found that not only did he not find any parasitized beetles in the field but even

when reared with known parasitoids of other coccinellids in the laboratory none of the

Rhyzobius larvae or adults were parasitized. This paralleled similar findings in nature

(Rubstov 1952, Smirnoff 1950). Stathas (2001) also found that even in the winter in

Greece R. lophanthae larvae could be found. In fact, both adults and larvae seem to be

able to remain active at temperatures as low as 8-90C (Stathas 2000, Cividanes and

Gutierrez 1996). Rhyzobius lophanthae also seems to be able to resist extreme heat,









because Atkinson (1983) found that adult R. lophanthae had an LD50 at about 420C.

Stathas (2000) found that in Greece R. lophanthae completed 6 generations a year, while

Smirnoff (1950) speculated that they may be able to complete as many as 7-8 generations

a year in Morocco.

This particular coccinellid was mentioned by Howard (1997) as a possible

biological control of CAS; but never mentioned again. Rhyzobius lophanthae has also

been labeled as "highly effective" in controlling CAS in Hawaii (Hara et al. 2004).

However, there are no research data to support this claim. In Tampa, Florida, adults and

larvae have been seen in large numbers feeding on CAS (personal observation 2004).

The fact that this beetle is already found in Florida makes it an appealing candidate for

the biological control of CAS. Label data with specimens in the FSCA indicate this

predator was captured feeding on Fiorinia theae Green, P. cockerelli, Pseudaulacaspis

pentagon (Targioni-Tozzetti), and Acutaspis morrisonorum Kosztarab.














CHAPTER 2
LIFE HISTORY OF Cybocephalus nipponicus ENDRODY-YOUNGA A PREDATOR
OF Aulacaspis yasumatsui TAKAGI (HOMOPTERA: DIASPIDIDAE)

Introduction

Cybocephalids are among the most economically important groups of natural

enemies against scale insects (Alvarez and Van Driesche 1998a). Larvae and adults are

voracious predators and have many desirable traits for use as biological control agents.

Cybocephalus nipponicus Endr6dy-Younga was released and later established in the

Washington D.C./Maryland area to combat the euonymus scale, Unaspis euonymi

(Comstock) (Drea and Carlson 1988, Drea and Hendrickson 1988). Alvarez and Van

Driesche (1998a) later released and established populations of C. nipponicus in New

England. This same species also was released, under the false identification of

Cybocephalus binotatus (Grouvelle), into the Miami area in 1998 for control of the cycad

aulacaspis scale, Aulacaspis yasumatsui Takagi (Anonymous 1998, Howard et al. 1999,

Howard and Weissling 1999). While C. nipponicus and C. binotatus appear similar, they

each have very distinctive male genitalia (Endr6dy-Younga 1971), and C. binotatus (Fig.

2-1A) has two large black spots on the pronotum, which are absent in C. nipponicus (Fig.

2-1B). Although C. nipponicus previously was established in Florida before 1998

(according to specimen label data in the Florida State Collection of Arthropods), its range

and abundance in the state before 1998 are unknown (Smith and Cave 2006b).

The cycad aulacaspis scale (CAS) is the most damaging scale found on Cycas in

Florida (Hodges et al. 2003). CAS is native to Thailand but is found throughout China









and southeastern Asia, as well as on several Caribbean islands, Florida, and Hawaii (Ben-

Dov et al. 2003). In 1992, CAS had become such a problem in Hong Kong that 70-100%

mortality was recorded in infested king sagos, Cycas revoluta Thunberg (Hodgson and

Martin 2001). The first detection of CAS in Florida occurred in 1996 in Miami at the

Montgomery Botanical Center. The scale was thought to have arrived on infested cycads

imported from southeastern Asia (Howard and Weissling 1999). By the end of 1997,

CAS had spread throughout Miami and as far north as Lake Okeechobee and could be

found on 20 species of cycads (Howard and Weissling 1999); however it seems to prefer

Cycas and Stangeria (Emshousen pers. comm. 2004). This led to the spread of CAS to

Hawaii in 1998 through the legal importation of infested cycads from Florida (Hodgson

and Martin 2001). At present, CAS has been reported from Pensacola east to

Jacksonville and south into the Florida Keys. Infested cycads in northern Florida were

suspected transplants from southern nurseries rather than natural progression of the scale

northward.

The objective of this study is to collect life history data on C. nipponicus using

CAS as prey, and compare these to the results from Tanaka and Inoue (1980) and Alvarez

and Van Driesche (1998a) using euonymus scale as prey. A better understanding of these

beetles using different prey will lead to greater understanding of how they perform as

biological control agents in the field, and consequently increase success in controlling

CAS.

Materials and Methods

A colony of CAS was reared on king sago in a sealed greenhouse to keep out

possible predators and/or parasitoids. Small king sago specimens were infested with

CAS by placing large numbers (>100) of eggs on each plant. Once infested, plants were









then placed in contact with other non-infested plants to spread non-parasitized scales to

other plants. Thus, we could be confident that parasitoids and predators did not invade

the colony.

A colony of C. nipponicus was initiated from individuals collected in south Miami

(N2538'21" W80020'09"). Aphanogmus albicoxalis Evans and Dessart (Hymenoptera:

Ceraphronidae) parasitizes the pupae of C. nipponicus in southern Florida (Evans et al.

2005), but this parasitoid was excluded by collecting only adult beetles and subsequently

rearing future generations in sealed cages (0.5m x 0.5m x 0.5m). Cycad leaves infested

with CAS were cut from the colony plant, with rachis bases placed in floral water tubes

for hydration, and then placed in rearing cages. The leaves and interior of the cage were

misted with distilled water every three days. A moist sponge also was provided for

hydration. Leaves were replaced every three weeks. Old leaves were held in separate

cages for three weeks to recover emerging beetles. Beetle and scale voucher specimens

were placed in the Florida State Collection of Arthropods (FSCA).

All life cycle studies were carried out in temperature- and humidity-controlled

cabinets set at 250 C with a relative humidity of 80% and a photoperiod of 14:10 (L:D).

Each treatment was initiated by isolating 25-30 mating pairs of beetles randomly selected

from the laboratory colony. Each pair was placed in a 25-dram plastic vial with one C.

revoluta leaflet infested with male and female CAS. After 24 hours the beetles were

removed and eggs collected from beneath the scale armor on the leaflet. Eggs were

measured (length and width), and then, to simulate natural conditions, placed on the

surface of a small, clean piece of C. revoluta leaflet and covered with the armor of an

adult female CAS. The leaflet piece was placed in the well of a rectangular tissue culture









dish, covered with parafilm, and placed in the environmental chamber. Eggs were

checked daily until larvae emerged.

Newly emerged larvae were placed in 25-dram plastic vials with freshly-cut, CAS-

infested leaflets on top of sterilized sand in the bottom of the vial. Old leaflets were

replaced with fresh, infested leaflets every five days throughout the life cycle. A fine

mesh cloth was placed over the top of the vials to allow airflow. Larval and pupal

development was checked daily. Upon emergence from the pupal case the adults

remained in plastic vials and were provided with 20-30 fresh CAS every 3 days. Adults

were checked daily until death.

Eggs, larvae, and pupae were critical-point dried in a Tousimis samdri-780 A,

and sputter coated with a gold-palladium alloy. Images were taken with a JEOL JSM-

5510LV scanning electron microscope and a Syncroscopy automontage photography

system.

Descriptive statistics were generated in SAS (2001) using a PROC UNIVARIATE

analysis. The dependent variable was number of days in each stage and the independent

variable was the stage itself. PROC UNIVARIATE and PROC t-test were used to

generate statistics for adult longevity.

Results and Discussion

The egg of C. nipponicus is elongate oval with both ends rounded, relatively large

measuring 0.42 mm by 0.20 mm (n=50) and usually light gray to purple. Eggs were

usually found singly inside the vacated tubular cover of a male scale or under the armor

of a female scale, usually with a live scale beneath but occasionally with a dead female.

During low scale density, as many as 5 eggs under one female armor were observed. An

egg deposited within the male scale cover fit snugly and had almost the same diameter as









the scale cover, thereby allowing only one egg to be placed in one male cover. The

surface was smooth aside from debris sticking to the surface (Fig. 2-2) and was slightly

tacky, allowing the egg to stick to the substrate. Females typically laid about 3 eggs per

day and on average 288 eggs in a lifetime (Alvarez and Van Driesche 1998a). Eggs

hatched about 7 days after oviposition (Table 2-1). Eyespots could be seen 1-2 days

before larval emergence.

When the larva emerged, the chorion split along the longitudinal axis and the larva

wriggled free. This process took about 15 minutes compared to the 30 to 45 minutes

reported by Blumberg and Swirski (1982) on the life history of Cybocephalus micans

Reitter and Cybocephalus nigriceps nigriceps (J. Sahlberg). The neonate larvae were

white or yellowish with long setae along the body, but after feeding for a day turned light

purple or lavender, with 4 black stemmata on each side of the head (Fig 2-3A, B). Not

only were larvae covered in long, slender setae but also shorter trumpet-shaped setae

(Fig. 2-4). After emergence, larvae immediately began to feed either on the scale eggs

sharing the space beneath the armor or rarely on the female scale. If an egg hatched in a

male scale cover, the larva would go to the nearest food source. Larvae continued to

move from scale to scale feeding on males, females, and eggs but spent the most time

underneath female armor. They also were seen cannibalizing other larvae when scale

density was extremely low, as mentioned by Alvarez and Van Driesche (1998a). Larvae

fed for 9 to 10 days.

Three instars (Fig. 2-5A, B) were observed, similar to C. micans and C. n.

nigriceps (Blumberg and Swirski 1982). However, Ahmad (1970) recorded four instars

in Cybocephalus semiflavus Champion. When molting, the cuticle ruptured along the top









of the head capsule and along the median dorsal region of the body. The larva emerged

from the anterior portion of the old exuviae first, and then wriggled vigorously to extract

the posterior body portion. The posterior portion of the old exuviae remained attached to

the substrate. Average larval development was about 14 days (Table 2-1).

If bright light was shone on larvae they immediately moved underneath a scale

armor or debris. Once disturbed larvae raised the head and body away from the leaf

surface, arching the body into a C-shape, and holding to the substrate with the conical

protuberance found on segments 8 and 9. This threat posture is similar to that used by the

larva to extract themselves from old exuviae.

Larvae became less active 2 to 3 days prior to pupation, stopped feeding, eventually

becoming immobile and attaching the posterior of the abdomen to the substrate before

forming the pupal case. Larvae gathered pieces of scale armor and incorporated these

pieces into an ovoid pupal chamber with one end flatter where it attached to the substrate

(Fig. 2-6A).

Pupal chambers often were found in the anterior portion between leaflet and leaf

rachis or near the leaflet base. However, pupal chambers also were observed along

leaflet and rachis. When not given access to scales, larvae used sand or other organic

material to construct the pupal chamber (Fig. 2-6B), sometimes attaching to the leaf or

dropping to the sand. Infrequently, larvae dropped to the sand and made a sand cocoon

even when scales were available. This behavior is common in Cybocephalus and has

been suggested or recorded for other species (Clausen and Berry 1932, Flanders 1934,

Smirnoff 1954, Blumberg and Swirski 1982). All pupae exhibited typical exarate









features (Fig. 2-7A, B). Pupation lasted about 18 days (Table 2-1). Beetles exited the

chamber by chewing an emergence hole.

There was no significant difference in developmental time from egg to adult

between sexes (t=1.50, df=52, P=0.1389). Total development from egg to adult lasted

about 40 days (Table 2-1), thus it is conceivable that 7-8 generations could be produced

per year in southern Florida or other areas with amenable temperatures. Alvarez and Van

Driesche (1998a) suggested that these beetles were capable of producing 3 generations

per year in New England.

Preoviposition lasted about 4 days (n=29) with some females laying eggs as early

as 2 days (n=2) after emergence. Adults (Fig. 2-8) began feeding soon after emergence

and consumed an average of 4 scales per day, with females typically eating more than the

males. Disproportionate feeding is probably due to size, because males are smaller than

the females. Maximum female longevity was 190 days, with an average of about 110

days (Table 2-1). Average longevity of males was 89 days (Table 2-1) with a maximum

of 155 days. Due to high variation, a t-test showed no significant difference between

longevity of the sexes (t=1.50, df=52, P=0.1389). These findings are intermediate

between the 78 days for males and 99 days for females reported by Alvarez and Van

Driesche (1998a) at 220 C and 122 days for males and 143 days for females reported by

Tanaka and Inoue (1980), who do not detail their experimental temperatures used.

Shorter male longevity may be due to their more active nature. Males were observed

moving around the cages more often than females. Often, 5 or 6 males would chase a

female for hours before finding a new female to pursue. The sex ratio of emerging adults

was 23:31 (male:female).









Our results were similar to those of Alvarez and Van Driesche (1998a) and Tanaka

and Inoue (1980). However, all stages developed slower in the former study (9.1 days for

eggs, 14.5 days for larvae, and 20.4 days for pupae) in comparison to our results, but this

maybe due to their lower experimental temperatures or the food source. Population

differences may account for some discrepancies in the life history parameters of the

beetles in each study. The beetles studied by Tanaka and Inoue (1980) originated in

Japan and those studied by Alvarez and Van Driesche (1998a) originated in Beijing,

China. Our beetles may have originated in Thailand, however, this is only speculation

considering the beetles were present in Florida before the recorded introduction in 1998

(Smith and Cave, in press). Mortality rates in our study also were slightly higher,

possibly due to high humidity. Fungal growth was a consistent problem in the humid

environment of our rearing chambers. Scales often became so encrusted on the cycad

leaflet that saprophytic fungi quickly spread. Alvarez and Van Driesche (1998a) did not

record relative humidity, thus a comparison cannot be made.

Mortality was highest during the larval stage. The first few days of larval

development proved to be most difficult (Fig. 2-9). Eggs and pupae seemed to be quite

hardy with mortality of 22% and 9%, respectively (Table 2-2), which are similar to the

14% and 8% mortality rates found by Alvarez and Van Driesche (1998a). Therefore, the

86% mortality rate in the larval stage (Table 2-2) is interesting and not predicted.

However, when reared on San Jose scale, Quadraspidiotusperniciosus (Comstock),

Alvarez and Van Driesche (1998a) found the larval stage of C. nipponicus had a

correspondingly high 77% mortality rate.









Certain aspects of the biology and life history of other Cybocephalus species have

been studied, typically as part of biological control projects, e.g., Cybocephalus rufifrons

Reitter (De Marzo 1995), Cybocephalusfreyi Endr6dy-Younga (Lupi 2003), and

Cybocephalusfodori Endr6dy-Younga (Katsoyannos 1984) have been studied in Europe;

and Cybocephalus semiflavus Champion (Ahmad 1970) and Cybocephalus gibbulus

Erichson (Nohara and Iwata 1988) have been studied in Asia. In the Middle East, several

studies have been carried out on C. aegyptiacus, C. binotatus, C. micans, and C. n.

nigriceps (Blumberg 1973, 1976; Blumberg and Swirski 1974 a,b, 1982). In Australia,

Kirejtshuk et al. (1997) described Cybocephalus aleyrodephagus and studied its life

cycle. The life histories of these species do not differ dramatically from C. nipponicus,

and there seems to be fairly consistent life cycle and feeding habits.

In the absence of prey, female cybocephalids are able to withhold eggs for up to 2

days, indicating that oviposition strategy is not only governed by food consumption but

also by qualitative features of the scale population (Alvarez and Van Driesche 1998b). In

the presence of high scale density, cybocephalids increase egg production until an

asymptote is reached. Again, this allows cybocephalids to maintain their populations at

low scale densities. If circumstances allow, female cybocephalids will lay only one egg

under a single scale cover. However, if the number of scales in a patch is low, and the

beetle cannot find new scales, it will lay eggs under a scale under which eggs are already

present (Alvarez and Van Driesche 1998b). The beetle larvae are forced to consume

more prey when feeding on younger scales, therefore for greater offspring survivability

female cybocephalids prefer oviposition on older scales that provide a larger food source

and require less searching (Alvarez and Van Driesche 1998b). Alvarez and Van Driesche









(1998b) found that the highest larval survival rate could be found in larvae feeding on

scales older than 30 days. Blumberg and Swirski (1982) noted that C. n. nigriceps rarely

laid eggs under dead female scales.

Evans et al. (2005) described two species ofAphanogmus that parasitize C.

nipponicus pupae. Aphanogmus inamicus Evans and Dessart emerged in quarantine from

hosts collected in Thailand, and A. albicoxalis occurs naturally in Florida. The latter

species is broadly distributed in southern Florida, with collection sites in Collier, Miami-

Dade, and St. Lucie counties. The levels of parasitism in C. nipponicus populations in

Florida are currently undetermined, but large numbers of these wasps occasionally have

been seen on cycads.

Cybocephalus nipponicus will probably always be considered a supplementary

predator for the control of CAS. However, some attributes make these beetles very

attractive as biological control agents, including a long lifespan, some ability to resist or

protect themselves from pesticides (Alvarez and Van Driesche 1998a, Katsoyannos 1984,

Kehat et al. 1974), and most importantly, the ability to persist at low scale densities

(Alvarez and Van Driesche 1998b). However, even in combination with the parasitoid

Coccobiusfulvus Compere and Annecke (Hodges et al. 2003) these beetles are unable to

adequately control populations of CAS in Florida.








































Figure 2-1. Dorsal habitus. A, Cybocephalus binotatus. B, Cybocephalus nipponicus.


Figure 2-2. Egg of Cybocephalus nipponicus.


I",;
.~ ~i :r~a, "

































Figure 2-3. First instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral
habitus.


Figure 2-4. Short trumpet-shaped setae on third instar larva of Cybocephalus nipponicus.

































Figure 2-5. Third instar larva of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral
oblique habitus.
























I'"


*,


i'


Figure 2-6. Pupal chambers of Cybocephalus nipponicus. A, Pupal chamber made from
female scale covers. B, Pupal chamber made from sand.


I'l if










































Figure 2-7. Pupa of Cybocephalus nipponicus. A, Dorsal habitus. B, Ventral habitus.


I ,





































Figure 2-8. Adult C. nipponicus, lateral habitus.


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Age in Days


Figure 2-9. Larval mortality over time.










Table 2-1. Development of Cybocephalus nipponicus on Aulacaspis yasumatsui.
Stage Mean (Days) SE Range (min-max) (Days) N
Egg 7.3 0.1 4(5-9) 131
Larva 13.7 0.1 4(12-16) 94
Pupa 18.6 0.2 6(16-22) 44
Total development time 39.5
Male life-span 89.1 9.7 146 (9-155) 23
Female life-span 110.0 9.6 175 (15-190) 31




Table 2-2. Mortality (%) of immature stages of Cybocephalus nipponicus during rearing
on Aulacaspis yasumatsui.
Stage (x) No. Alive at Start (lx) No. Dying in Stage (dx) Apparent Mortality (qx)
Egg 200 44 0.22
Larva 156 134 0.86
Pupa 22 2 0.09
Adult 20














CHAPTER 3
PESTICIDE SUSCEPTIBILITY OF Cybocephalus nipponicus AND Rhyzobius
lophanthae (COLEOPTERA: CYBOCEPHALIDAE, COCCINELLIDAE)

Introduction

Beetles of the families Coccinellidae and Cybocephalidae are the most

economically important groups of predators of diaspidid scales in the world (Blumberg

and Swirski 1982). Cybocephalus nipponicus Endrody-Younga (Cybocephalidae) and

Rhyzobius lophanthae Blaisdell (Coccinellidae) are commonly used as biological control

agents for many armored scale pests. Rhyzobius lophanthae has been established in

Florida since the 1930s (according to specimen label data in the Florida State Collection

of Arthropods). Cybocephalus nipponicus, misidentified as Cybocephalus binotatus

Grouvelle, was recently released in south Florida in an effort to control the cycad

aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Anon. 1998; Howard et al. 1999;

Howard and Weissling 1999). CAS is the most economically damaging scale to cycads

that the state of Florida has ever seen (Hodges et al. 2003). Although C. nipponicus is

present in Hawaii (Heu and Chun 2000), R. lophanthae is usually suggested as the better

control agent of CAS (Heu et al. 2003; A. Hara, personal communication). In both

places, CAS has continued to spread and multiply. A more promising approach to

controlling CAS would be one using integrated pest management (IPM). In this manner,

a combination of pesticides and biological control would be used to combat CAS.

There has been some success controlling CAS with various pesticides. Oils, either

an ultra-fine horticultural oil or a product containing fish oils, seem to be the most









effective chemical control method (Hodges et al. 2003). This is not surprising given that

oils have long been used to control armored scale insects. The oil not only covers the

insects and suffocates them but also covers the surface of the plant making it difficult for

crawlers to settle onto the plant (Howard and Weissling 1999). Soaps are quite popular

with homeowners; but they must be applied frequently, in some cases once a week

(personal observation). The effective application of pesticides for control of CAS is

difficult due to the scale's tendency to heavily infest the abaxial surface of leaves, which

is difficult to spray (Howard and Weissling 1999). In the case of C. revoluta, the

architecture of the plant itself, with the margins of the leaflets curling down and inward to

form an arch on the abaxial surface of the leaflet, makes foliar treatments inefficient

(Hodges et al. 2003). Frequent or "as needed" applications of oils seems to be the most

effective technique for controlling CAS, and by mixing oil with contact pesticides such as

malathion, even greater scale mortality can be achieved (Hodges et al. 2003). The use of

systemic pesticides such as dimethoate and contact pesticides like methidathion has

yielded mixed results, being very effective in some instances and completely ineffective

in other cases (Hodges et al. 2003). Imidacloprid used as a soil drench can be very

effective, but Howard and Weissling (1999) found that this product had to be mixed at

very high concentrations to be effective. This product can also be used as a foliar spray.

The reproductive biology of C. nipponicus makes it a good biological control

agent. Alvarez and Van Driesche (1998a) found that, at low scale densities, C.

nipponicus was able to maintain its populations and maintain populations of euonymus

scale, Unaspis euonymi (Comstock), and San Jose scale, Quadraspidiotusperniciosus

(Comstock), in check. In the presence of greater scale densities, C. nipponicus will









increase its egg production accordingly. With a total life cycle from egg to adult only

taking around 44 days (Smith and Cave 2006b), it is conceivable that 5-6 generations

could be produced every year in Florida. Cybocephalus nipponicus is available

commercially in the U.S. market.

Rhyzobius lophanthae is an exceptional biological control agent because of its high

fecundity, lack of parasitoids, the absence of diapause, and resistance to low temperatures

especially in the immature stages (Rubstov 1952; Smirnoff 1950; Stathas 2000). Female

R. lophanthae are able to lay hundreds of eggs in a lifetime (Stathas 2000). Rhyzobius

lophanthae also seems to be able to resist extreme heat. Atkinson (1983) found that adult

R. lophanthae could not survive for long at 420 C. Rhyzobius lophanthae is also available

commercially in the U.S. market.

This study was conducted to determine the susceptibility of C. nipponicus and R.

lophanthae to six pesticides commonly used in the control of CAS. Given the established

presence of both predators on cycads in south Florida and their commercial availability, it

is very important to learn what effects the commonly used pesticides against CAS will

have on them. This information is vital for development of IPM programs aimed at

controlling CAS.

Materials and Methods

Insects

Adult R. lophanthae were reared at and purchased from Rincon-Vitova Insectaries

(Ventura, California). Adult C. nipponicus were also purchased from Rincon-Vitova but

were reared by Philip Alampi Beneficial Insect Laboratory, New Jersey Department of

Agriculture. Both beetle species were maintained in Plexiglas cages at 250 C prior to

testing. All life stages of CAS were provided as a food source.









Food was not provided during testing because of the very small size of the beetles

(1 mm and 2.5 mm in length). The beetles could have conceivably perched on the food

source for long periods of time, never coming into contact with the walls of the treated

vial. Preliminary studies indicated that a 24-hour period without food would not unduly

stress the beetles. On average, untreated C. nipponicus survived for 8-9 days (n=30) and

untreated R. lophanthae lived for 5-6 days (n=30) before dying of starvation. Cotton used

to stopper the vials was soaked in water to prevent dehydration.

Bioassays Using Coated Glass Vial Method

A coated glass vial method (Plapp 1971; Amalin et al. 2000; Snodgrass 1996;

Snodgrass et al. 2005) was used to determine the chemical susceptibility of adult R.

lophanthae and C. nipponicus to six pesticides used to control CAS (Howard et al. 1997;

Howard and Weissling 1999; Weissling et al. 1999; Hodges et al. 2003; Emshousen and

Mannion 2004). This is a very effective method for testing the chemical susceptibility of

small arthropods (Amalin et al. 2000) such as R. lophanthae and especially C. nipponicus

because of its extremely small size. The six pesticides tested were fish oil emulsion

(Organocide), insecticidal soap (Garden Safe, Inc.), imidacloprid (Provado),

malathion (Spectracide, Inc.), methidathion (Supracide), and dimethoate (Cygon).

The fish oil and insecticidal soap were purchased at commercial grade, while the

imidacloprid (99% purity), malathion (98% purity), methidathion (98.6% purity), and

dimethoate (98.7% purity) were purchased in the technical grade from Chem Service

(West Chester, PA).

All pesticides were dissolved in acetone, except the insecticidal soap which does

not dissolve in acetone. Instead, the insecticidal soap was dissolved in 95% ethanol. The

fish oil was shaken in a paint shaker after being placed in acetone in order to break up the









oil into fine globules. Each pesticide was separated into three dilutions: field rate, twice

field rate, and one-half field rate. The field rate was taken from label data for each

pesticide as directed for use against scale insects. A small amount (0.5 ml) of the

pesticide working solution was dispensed into 20-ml scintillation vials. Concentrations

of active ingredient for the working solution and the amount of active ingredient residue

within the vials can be seen in Table 3-1. Vials were hand turned until the acetone or

ethanol completely evaporated leaving an insecticidal residue on the inner surface. Vials

treated with only acetone or ethanol, as well as untreated vials, were used as controls. A

single beetle was placed into a treated vial. All beetles had emerged from pupae within

the previous 14 days. Vials were sealed with cotton soaked in water allowing the beetles

to drink. Vials were placed upright in a ventilated cabinet with a fume hood and at a

constant temperature of 250 C and 80% relative humidity for 24 hours. For each treatment

of 10 beetles, 5 females and 5 males were used. Each treatment of 10 beetles was

replicated 3 times for each dosage. All trials were carried out the same day as the

pesticide was applied to the vials.

Mortality of beetles was determined immediately after the 24-hour period. A beetle

was considered dead if it was not moving or could not right itself. Percent survivorship

was measured as the proportion of 30 beetles alive after a 24-hour exposure to the

pesticides.

Statistical Analysis

All descriptive statistics were generated in EXCEL (Microsoft 2000). The

mortality rates for each pesticide were compared using the Student-Newman-Keuls mean

separation test (SAS Institute 2001).









Results

Of the six pesticides tested on adult C. nipponicus and R. lophanthae, three

(methidathion, dimethoate, and malathion) caused >90% mortality at all concentrations,

while the other three (fish oil, insecticidal soap, and imidacloprid) were less toxic but still

caused very high mortality (Tables 3-2 & 3-3).

Effects of Pesticide on C. nipponicus

Cybocephalus nipponicus was extremely susceptible to all pesticides. The three

least toxic pesticides were imidacloprid, insecticidal soap, and fish oil (Table 3-2). There

were significant differences in mortality between concentrations amongst these three

pesticides (Table 3-4). Fish oil was still quite toxic to this beetle, and due to its very

small size, C. nipponicus would often get trapped in small globules of oil, eventually

dying from suffocation.

Effects of Pesticide on R. lophanthae

Rhyzobius lophanthae was more tolerant than C. nipponicus to the experimental

pesticides, although mortality rates were also high for this species. The three least toxic

pesticides to R. lophanthae were imidacloprid, insecticidal soap, and fish oil (Table 3-3).

There were significant differences in survivorship between concentrations of these three

pesticides (Table 3-4). Rhyzobius lophanthae, about twice the size of C. nipponicus,

had much less difficulty traversing oil globules on the surface of the vials.

Discussion

There was a significant difference between mortality in the control and that of even

the lowest pesticide concentration. This sensitivity to pesticides makes an IPM approach

to the control of CAS quite difficult. Unfortunately, most of the success in chemically









controlling CAS has involved very toxic pesticides often being used at higher than

recommended doses (Howard and Weissling 1999, Weissling et al. 1999).

The high mortalities experienced by C. nipponicus and R. lophanthae are not

unexpected. Nakao et al. (1985) found that all 18 species of Coccinellidae inhabiting

Japanese citrus groves were severely affected by the application of pesticides, including

methidathion and dimethoate. They also found that Cybocephalus gibbulus Erichson,

one of the most common scale predators found in Japanese citrus groves, was virtually

eliminated by long-term pesticide use. Oils have proven to be the most effective

pesticides used against many plant-sucking pests, while maintaining the natural enemy

populations. Erkilic and Uygun (1997) found that oils were much less toxic to

Cybocephalusfodori minor (Endrody-Younga) and Chilocorus bipustulatus (Linnaeus)

than was methidathion. In fact, they went as far as to say that methidathion should not be

used in IPM programs.

In natural conditions, the predatory beetles may not likely be in contact with the

pesticide for as long as the exposures in this experiment. However, C. nipponicus and R.

lophanthae are uniquely suited for life in chemically-treated environments. Both beetle

species place their eggs underneath the scale cover and at least part of larval development

takes place beneath the armored scale, allowing the beetles some protection from both the

elements and pesticides (Smirnoff 1950; Alvarez and Van Driesche 1998; Stathas 2001).

In Greece, Katsoyannos (1984) found that C. fodori was able to survive in pesticide-

treated fruit orchards. In date palm plantations in Israel, Kehat et al. (1974) found that,

while all coccinellids in a chemically-treated plantation died, species of Cybocephalus

survived.









It is apparent from these tests that, for some pesticides, the lower the concentration

of the pesticide the higher the survivorship. However, these tests were conducted in a

laboratory environment in which the test subjects were in constant contact with the

pesticide for 24 hours. A whole host of factors, such as humidity, UV degradation,

evaporation, and precipitation, will influence pesticide activity in the field. Nevertheless,

whenever possible, insecticidal soaps and fish oils should be used. While many

homeowners use various types of soaps to treat CAS, this method requires treatment

every 10 to 14 days, thus increasing exposure of the beetles to the pesticide. If more toxic

pesticides must be used, then applying them to "hot spots" rather than broadcast spraying

may protect the scale predators from complete annihilation. This type of selective

spraying may also protect other entomophagous insect populations from being decimated

(Kuznetsov 1997). The results of these laboratory experiments yield some baseline data

from which more research in the field can be conducted.










Table 3-1. Field rates (IX) for each pesticide used.
Insecticide Working Solution (pg*AI/ml)
Organocide 47000
Insecticidal Soap 512300
Imidacloprid 106
Methidathion 233
Dimethoate 305
Malathion 1990


Insecticide residue (pg*AI/cm2)
8.29
27.71
2.40
5.26
6.91
45.07


* Al = Active Ingredient


Table 3-2. Percent mortality of Cybocephalus nipponicus per 30 individuals exposed. X
field rate.
% beetle % beetle % beetle % beetle
Pesticide mortality mortality mortality mortality


at OX


Organocide
Insecticidal Soap
Imidacloprid
Methidathion
Dimethoate
Malathion
Control (Acetone)
Control (Ethanol)
Control (No coating)


at 0.5X
83
66
76


at 1X
100
86
93
100
96
100


at 2X
96
96
100
100
100
100


Table 3-3. Percent mortality ofRhyzobius lophanthae per 30 individuals exposed. X
field rate.


Pesticide


% beetle
mortality
at OX


% beetle
mortality
at 0.5X


Organocide
Insecticidal Soap
Imidacloprid
Methidathion
Dimethoate
Malathion
Control (Acetone)
Control (Ethanol)
Control (No coating)


% beetle
mortality
at 1X
83
76
80
100
96


% beetle
mortality
at 2X
100
96
100
100
100







41


Table 3-4. Student-Newman-Keuls test showing ranked values of mortality of adult
Cybocephalus nipponicus and Rhyzobius lophanthae.
C. nipponicus Dose Imidacloprid Organocide Insecticidal soap
O.OX 2.0A 2.0A 2.0A
0.5X 5.5B 5.3B 5.3B
1.0X 8.5C 8.6C 8.3C
2.0X 10.0C 10.0C 10.3C

R. lophanthae Dose Imidacloprid Organocide Insecticidal soap
0.OX 2.0A 2.0A 2.0A
0.5X 5.6B 5.0B 5.3B
1.0X 7.3B 8.0C 7.8C
2.0X 11.0C 11.0D 10.8D
Mean ranks within columns with the same letter are not significantly different (P=0.05).














CHAPTER 4
THE CYBOCEPHALIDAE (COLEOPTERA) OF AMERICA NORTH OF MEXICO

Introduction

The family Cybocephalidae consists of seven genera: Cybocephalus (Erichson

1844), Endrodiellus (Endrddy-Younga 1962a), Hierronius (Endrddy-Younga 1968),

Horadion (Endrddy-Younga 1976), Pastillodes (Endrddy-Younga 1968), Pastillus

(Endrddy-Younga 1962a), and Pycnocephalus (Sharp 1891). By far the largest of these

is Cybocephalus, which contains more than 150 described species found throughout the

world (Tian 2000, Yu and Tian 1995).

The Cybocephalidae differ in many ways from the Nitidulidae. They are predatory,

almost exclusively feeding on scale insects, while nitidulids are known for feeding on

decaying plant material and fruits, plant sap, fungi, and occasionally pollen and honey.

The morphology of the Cybocephalidae is also quite different from that of the

Nitidulidae. Cybocephalid adults have a 4-4-4 tarsal formula instead of 5-5-5 found in

Nitidulidae. There are 5 visible ventral plates (leaving out the male anal plate) and 5

abdominal spiracles in cybocephalids instead of the 6 and 6 that occur in nititdulids. The

body of cybocephalids is retractile allowing the mandibles in repose to rest against the

metasternum, unlike any other nitidulid. The larvae of Cybocephalidae have a head

without dorsal sutures, lack pregomphi and urogomphi on abdominal tergite XI, and have

hypostomal rods with divergent hypostomal ridges present posteriorly, hypopharynx

without a sclerome and bracons, maxillae without mola, and annular spiracles with 2

lateral air tubes. In contrast, the larvae of Nitidulidae have pregomphi and urogomphi, no









hypostomal rods but with hypostomal ridges strongly convergent posteriorly,

hypopharynx with a sclerome and bracons, maxillae with raised mola, and biforous

spiracles (Kirejtchuk 1997). These larval differences were also illustrated by Boving and

Craighead (1931) and Hayashi (1978). In spite of this, the group has been shuffled

between family (Jacquelin du Val 1858, Murray 1864, Boving and Craighead 1931,

Parsons 1943, Smirnoff 1954, Endrody-Younga 1968) and subfamily (Erichson 1844,

Horn 1879, Grouvelle 1913, Kirejtshuk 1997, Habeck 2002) status. We chose to

recognize the Cybocephalidae as a family in accordance with the work of the world

expert in this group, Sebastian Endrody-Younga.

The adults of Cybocephalus are often confused with those of Clambidae,

Phalacridae, and sometimes Coccinellidae (Microweisea Cockerell and Guiiithiil eil'e

Gordon). The major feature that distinguishes members of Cybocephalus from all of

these families is their extremely broad head. The head of Cybocephalus is almost as wide

as the pronotum, unlike that of all the aforementioned families. Adult clambids have

extremely enlarged hind coxae, a clypeus that entirely covers the mouthparts, and a 2-

segmented antennal club, whereas cybocephalids have small hind coxae, a relatively

short clypeus and a 3-segmented antennal club. Phalacrids have a much more elongate

club than cybocephalids and a 5-5-5 tarsal formula rather than the 4-4-4 formula found in

cybocephalids. Gihiuthii, eI'et has an extremely long head compared to a short, wide

head in cybocephalids. Also, both Giiithihi eilea and Microweisea have the head

inserted into the prothorax, unlike cybocephalids which have the head completely outside

the pronotum, and both genera have distinctive tarsomeres easily distinguishing them

from the Cybocephalidae.









Cybocephalids are primarily known for feeding on armored scales (Diaspididae)

(Flanders 1934, Clausen 1940, Vinson 1959, Endrody-Younga 1968, Blumberg and

Swirski 1974a,b, Rosen and DeBach 1978, Alvarez and Van Dreische 1998a). However,

they have also been reported feeding on whiteflies (Aleyrodidae) (Clausen and Berry

1932, Chandra and Avasthy 1978, Swirski, et al. 1987, Kapadia and Puri 1993, Kajita et

al. 1991, Kirejtshuk et al. 1997, Ramani 2000, Tian and Ramani 2003), mealybugs

(Pseudococcidae) (Endrody-Younga 1982), and citrus red mite, Panonychus citri

(McGregor) (Tanaka and Inoue 1980).

These minute beetles can be found throughout the world, but relatively few of them

have been studied. Grouvelle (1913) described 5 species of Cybocephalus from the

Seychelles, while Vinson (1959) found at least 8 new species in addition to the two

known species of Cybocephalus from the small Mascarene Islands. More recently, Tian

and Peng (1997) recorded at least 8 species of Cybocephalus from Hainan Island in

China. It seems very unlikely that all of these island groups, with such a finite amount of

land area, would contain such rich Cybocephalus diversity while larger land masses

would not. Considering that there are 150 described species of Cybocephalus, these

studies would seem to indicate that these beetles are often overlooked or simply ignored.

Cybocephalus species are particularly difficult to identify because they are very

small (0.5-2.5 mm.), compact, extremely convex and have a deflexed head and the ability

to retract their appendages. For this reason, descriptions of male genitalia are an

important accompaniment to any description or key to species within this genus (Vinson

1959). Male genitalia in conjunction with other useful diagnostic characters such as the

form of the antenna can make identification less difficult.









The objectives of this study are to 1) develop techniques for the identification of

the cybocephalids of America north of Mexico; 2) validate and review the identification

of adventive cybocephalids released in Florida; 3) describe new North American species;

4) determine major prey species of these cybocephalids; and 5) report the known

distribution of each species in America north of Mexico.

Because some species of Cybocephalus are being released in several countries as

biological control agents, it is absolutely imperative that a clear and concise form of

identification be devised. "Taxonomic support is especially important because correct

identification of pests and their natural enemies are [sic] absolutely essential for effective

biological control" (Van Driesche and Bellows 1993).

Taxonomic History

The genus Cybocephalus was described by Erichson in 1844, who also described

five species from Europe, Africa, and Asia. LeConte (1863) described the first New

World cybocephalid, Cybocephalus nigritulus. Horn described a second species,

Cybocephalus californicus, from North America in 1879. Since that time no new species

of Cybocephalus have been described from America north of Mexico. The only other

species known to occur in this region is Cybocephalus nipponicus Endrody-Younga,

which was brought from Asia as a biological control agent in the late 1980s (Drea and

Carlson 1988). Most taxonomic research on this group was carried out by Endrody-

Younga (1962a,b, 1963, 1964a,b, 1965, 1967a,b, 1968, 1969, 1971a,b, 1976, 1979, 1982,

1984) on beetles found in Africa, Eurasia, and Oceania. Lately, an enormous amount of

work has been done on the cybocephalids of China and southern Asia by M. Tian and co-

workers (Tian 1995, 1996, 2000; Tian and Pang 1994; Tian and Ramani 2003; Tian and

Yu 1994; Tian and Peng 1997; Yu, 1994, 1995a,b; Yu and Tian 1995).









Materials and Methods

Materials

For this study, 871 specimens belonging to the genus Cybocephalus were

examined. The holotype of C. nigritulus, the lectotype of C. californicus and a topotype

of Cybocephalus binotatus (Grouvelle) were included. Specimens were borrowed from

the following institutions and private collections; names of the curators and owners are in

parentheses:

AAIC Albert Allen Insect Collection, Boise, ID, (Albert Allen)

BMNH Natural History Museum, London [formerly British Museum (Natural History)],
UK, (Maxwell Barclay)

CSCA California State Collection of Arthropods, Sacramento, CA, (Chuck Bellamy)

CNCI Canadian National Collection of Insects, Ottawa, Ontario, (Anthony Davies)

EMEC Essig Museum of Entomology, University of California, Berkeley, CA, (Cheryl
Barr)

FSCA Florida State Collection of Arthropods, Gainesville, FL, (Paul Skelley and
Michael Thomas)

HGIC Holly Glenn Insect Collection, Homestead, FL, (Holly Glenn)

INHS Illinois Natural History Survey, University of Illinois, Champaign, IL, (Colin
Favret)

LACM Los Angeles County Museum of Natural History, University of California, Los
Angeles, CA, (Weiping Xie)

LSAM Louisiana State Arthropod Museum, Louisiana State University, Baton Rouge,
LA, (Andrew Cline and Chris Carlton)

MTEC Montana Entomology Collection, Montana State University, Bozeman, MT,
(Michael Ivie)

MCZC Museum of Comparative Zoology Collection, Harvard University, Cambridge,
MA, (Philip Perkins)









OSUC Ohio State University Collection, Museum of Biological Diversity, Columbus,
OH, (Norman Johnson)

OSAC Oregon State Arthropod Collection, Oregon State University, Corvallis, OR,
(Jason Leathers)

SBMN Santa Barbara Museum of Natural History, Santa Barbara, CA, (Michael
Caterino)

SEMC Snow Entomological Museum, University of Kansas, Lawrence, KA, (Zack H.
Falin)

TAMU Texas A&M University, Department of Entomology, College Station, TX, (Ed
G. Riley)

TRSC Trevor Randall Smith Collection, Gainesville, FL, (Trevor Randall Smith)

UCDC University of California Davis, R. M. Bohart Museum of Entomology, Davis,
CA, (Steve L. Heydon)

UCRC University of California Riverside, Entomology Research Museum, Riverside,
CA, (Doug Yanega)

UCFC University of Central Florida Insect Collection, University of Central Florida,
Orlando, FL, (Stuart Fullerton)

WFBM W. F. Barr Entomological Museum, University of Idaho, Moscow, ID, (Frank
Merickel)

USNM United States National Museum, Smithsonian Institute, Washington D.C., (Gary
Hevel)

Methods

If possible, genitalia were removed using minute pins glued to wooden applicator

sticks. The minutes were bent and twisted into whatever shape tools were necessary.

While it is possible to remove only the genital plate and then extract male genitalia, this

technique is extremely difficult and time-consuming. All dissections took place in

glycerine due to the convex body form of Cybocephalus beetles. Typically, the abdomen

was separated from the rest of the body and the entire aedeagus (Fig. 4-1) removed from

the abdomen. This technique leaves most of the specimen intact, and the removed









abdomen can be glued to the point behind the specimen. The tegmen and median lobe

are compressed and held together by the tegminal (lateral) struts. To separate these two

parts, a minute was wedged between the median lobe and the tegminal strut breaking

one lobe of the tegminal strut, and separating the two pieces. In some cases, the internal

sac can be removed, and the tegmen can be moved backwards sliding both tegminal struts

over the posterior portion of the median lobe as well as the median strut, effectively

separating the two parts without damaging the tegminal struts, the median strut, or the

dorsal piece of the tegmen. The median lobe and tegmen (basal plate) can then be slide-

mounted or, to avoid distortion, mounted on a point in dimethyl hydantoin formaldehyde

(DMHF). This solution is water soluble and dries clear. If possible, genitalia should be

mounted on very shallow depression slides to avoid distortion (Fig. 6, 19, 32). This is

especially true of a median lobe with a large and raised median plate.

Specimens were cleared in 10% KOH at 24C in preparation for disarticulation.

After 24 hours, beetles were sufficiently cleared and softened for dissection. Due to their

convex body form and very small size, specimens were disarticulated in glycerine. All

disarticulated parts, including genitalia, were then washed in 95% ethanol and mounted

on microscope slides using Hoyer's solution. Label data were copied onto slides

verbatim with label breaks indicated by a slash (/).

Definitions

Median lobe (Fig. 4-1): Also referred to as penis (Endr6dy-Younga 1968, 1971a, 1971b;

Kirejtshuk et al. 1997; Lupi 2003; Yu 1995a, 1995b).

Basal plate (Fig. 4-1): This is a reference to the basal plate of the tegmen (Endrody-

Younga 1968, 1971a, 1971b; Lupi 2003; Yu 1995a, 1995b).









Cybocephalus Erichson 1844

Cybocephalus Erichson 1844: 441-442.

Redescription. Form: Ovate and very convex; body contractile (Fig. 4-2). Head:

Broad and deflexed. Labrum emarginate. Epistoma slightly prolonged at middle.

Mandibles in repose resting against metasternum, acute at tip with a small tooth

posteriorly. Maxillae with one lobe. Antennae slightly longer than width of head,

antennal club flat with 3 antennomeres, antennal grooves small and convergent. Scape

large and round. Thorax: Pronotum margined at base, covering base of elytra, sides very

short. Prostemum acutely carinate in front, not prolonged behind the procoxae, procoxal

cavities open behind. Mesostemum broad, oblique. Metasternum protuberant and

clothed in hairs. Both the meso- and metasternum are impressed for the reception of the

middle and hind legs. Scutellum: Large, triangular. Elytra: Covering or nearly covering

tip of the abdomen, apices curved. Abdomen: Five visible ventral plates (omitting the

small male anal plate), and 5 abdominal spiracles. Legs: Tibiae simple, tarsi four-

jointed, each tarsomere slightly dilated ventrally, second and third tarsomeres bilobed,

claws simple. Median lobe: Trunk heavily sclerotized and dorsoventrally compressed.

Tegmen: No parameres.

Key to the species of Cybocephalus of America north of Mexico

1. Antennal club without a serrated margin (Fig. 4-12); scutellum with slightly
concave margins (Fig. 4-11) ........ .. ................................ kathrynae new species

1'. Antennal club with a serrated margin (Fig. 4-4); scutellum with straight or slightly
convex m argins (Fig. 4-10)........................................................... ............... 2

2(1') Terminal antennomere of the antennal club rounded (Fig. 4-30)..........................
...... .. .......... ............................... ..............randalli new species

2' Terminal antennomere of the antennal club truncate (Fig. 4-17) or slightly
em arginate (Fig. 4-3, 4-4) .............................................. ...... .............










3(2'). Antennomere 3 slightly shorter than 4 and 5 combined; male bicolored with head,
prothorax, and mesosternum yellow or tan, rest of body black (Fig. 4-29); basal
plate coming to a rounded point (Fig. 4-26); median lobe as in Figs. 4-24, 4-25 .....
......... ....................................... .......... ....... .. nipponicus Endrody-Y ounga

3'. Antennomere 3 as long or longer than 4 and 5 combined; male with head,
prothorax and mesothorax black or dark brown ............................................4

4(3'). All legs and antennae brown or black; elytral apices with a large and wide
impunctate area of yellowish translucence; male basal plate evenly rounded,
without a protuberance (Fig. 4-8); median lobe as in Figs. 4-5-4-7 ...................
..................................................................................... californicus H orn

4'. At least profemora yellow or amber and usually antennae amber or tan;
translucent impunctate area at the apices of elytra much smaller; male basal plate
with a protuberance in the middle (Fig. 4-21) median lobe as in Figs. 4-18-420......
................... ...... .................................... ........ .... .......... ...... .. n ig ritu lus L eC onte

Cybocephalus californicus Horn

(Figs. 4-3-4-9)

Cybocephalus californicus Horn 1879: 320-321.

Diagnosis. Male and female are black, brown, or aeneous. Antennal club is smaller

than the eye and truncate or slightly emarginate at the apex, unlike the 11th antennomere

of C. randalli, which is rounded. Each antennomere of the club is distinctly separated,

forming a serrated margin, unlike C. kathrynae which has a smooth club margin. Apices

of the elytra have a large, wide area of yellowish translucence that is without punctation,

which distinguishes this species from C. nigritulus. In males, the basal plate and median

lobe are easily distinguished from those of all other species.

Redescription. Male. Form: Elongate oval; contractile; strongly convex

dorsally. Length: 0.95-1.30 mm (measured from apex of clypeus to apex of elytra);

breadth: 0.85-1.20 mm (measured at base of elytra). Color: Head, thorax, elytra and

underside black with surface sometimes aeneous, lateral margin of pronotum and









posterior margin of elytra yellowish and translucent; legs and antennae brown or black.

Head: Broad and convex, clypeus moderately produced, narrow, and slightly reflexed.

Eyes large, oblong, with internal margins distinct. Genae not visible from above but

slightly explanate when viewed laterally. Dorsal surface smooth under high

magnification, distinctly alutaceous and finely punctate. Antennae 11-segmented

including a club with 3 antennomeres, club length about 12-2/3 size of the eye. Club flat

and distinctly separated from funicle and with a distinctly serrated margin. First club

antennomere wider than long, second club antennomere larger than either first or third

club antennomere and about as long as wide. Terminal club antennomere truncate (Fig.

4-3) or emarginate (Fig. 4-4), setose and about as long as wide. Antennomere 3 as long

or slightly longer than antennomeres 4 and 5 combined. Pronotum: Strongly convex,

lateral margins curved; anterior angle more narrowly arcuate than posterior. Surface

distinctly alutaceous with fine punctation. Scutellum: Alutaceous and triangular with

straight to slightly convex margins. Elytra: Uniform width narrowing at apical 1/5.

Strongly convex, sides slightly sinuous and apices rounded. Length slightly shorter than

combined width (30:38). Uniformly punctate along dorsal surface, smooth at sides and

base with a large impunctate area at apices of elytra. Appearing alutaceous under high

magnification. Median margin and apices of elytra bordered. Underside: Metasternum

alutaceous, roughly punctured, and clothed in long coarse hairs. Abdominal sternites

alutaceous and punctate with long coarse hairs thinly covering the surface. Legs:

Femora glossy, broad, flattened and sparsely covered with short hairs. Pro- and

mesofemora about the same width throughout length. All tibiae slightly but distinctly

curved and dilated towards apex. The protibiae with short hairs along outer margin.









Meso- and metatibiae with long stiff hairs along outer margin. Four tarsomeres, claw

tarsomere as long or almost as long as 2 tarsomeres preceding it. Median lobe: Sides

parallel or slightly divergent curving into a point with declivous sides from apex (Figs. 4-

5, 4-6). In profile, strongly curved from middle (Fig. 4-7). Median plate on surface

elevated. Basal plate: Sides parallel at base, evenly rounded at apex (Fig. 4-8).

Female. Nearly identical to male.

Geographic distribution. Arizona, British Columbia, California, Colorado, Idaho,

Montana, Nevada, New Mexico, Oklahoma, Oregon, Texas, Utah, Washington (Fig. 4-9).

Hosts. This species is known to feed on or was found in association with Aonidiella

aurantii (Maskell) (from label data), Chionaspispinifolia (Fitch) (from label data),

Diaspidiotus perniciosus (Comstock) (Heintz 2001), Diaspis echinocacti (Bouche) (from

label data), Diaspis manzanitae Whitney (from label data), Ehrhornia cypressi (Ehrhorn)

(Flanders 1934; Kartman 1946; Clausen 1940), Lecanium corni Bouche (Heintz 2001),

Lepidosaphes beckii (Newman) (from label data), Mercetaspis halli (Green) (Kartman

1946), and Parlatoria blanchardi Targioni Tozzetti (from label data), all of which belong

to the family Diaspididae. It has also been seen in association with Phoenicococcus

marlatti Cockerell (from label data) belonging to the family Phoenicococcidae, and some

species of aleyrodids (from label data).

Type material examined. The so-called lectotype in the MCZC is a male

specimen glued to a point with the following labels: Cal (printed) [small white

rectangular label with the edge of one side dipped in purple ink] / LectoTYPE (printed)

3216 (handwritten) [red rectangular label] / Horn Coll H (printed) 3751 (handwritten)

[white rectangular label] / C. californicus Cr. (handwritten) [white rectangular label] /









MCZ Type (printed) 7970 (handwritten) [red rectangular label] / Cybocephalus

californicus Horn Det: Trevor Smith (printed) [white rectangular label]. While this

specimen is a syntype, it is not a valid lectotype because this designation was never

published. This is consistent with many of Horn's type specimens that had lectotype

labels placed on them around 1930 without corresponding publications. This occurred

when the Horn collection was located at the Academy of Natural Sciences, Philadelphia

(Philip D. Perkins 2005, personal communication). Two female specimens, labeled as

paratypes in the MCZC, were also examined. The first "paratype" has the following

labels: Cala (printed) F (handwritten) [small white rectangular label] / Para-Type (printed)

3216.4 (handwritten) [blue rectangular label] / Horn Coll H (printed) 3751 (handwritten)

[white rectangular label] / Type 7970 (handwritten) [red rectangular label] /

Cybocephalus californicus Horn Det: Trevor Smith (printed) [white rectangular label].

The second paratype label has the following labels: Cal (printed) j (handwritten) [small

white rectangular label] / Para-Type (printed) 3216.3 (handwritten) [blue rectangular

label] / Horn Coll H (printed) 3751 (handwritten) [white rectangular label] / Type 7970

(handwritten) [red rectangular label] / Cybocephalus californicus Horn Det: Trevor Smith

(printed) [white rectangular label]. Two male syntypes in the MCZC were also

examined. The syntypes are both glued to a single card with the following labels: Type

(printed) 7970 (handwritten) [orange rectangular label] / Cybocephalus californicus Horn

(handwritten with Cr. crossed and Horn written in a different colored ink) [large white

rectangular label] / Cybocephalus californicus Horn Det: Trevor Smith (printed) [white

rectangular label].









Other material examined. CANADA: BRITISH COLUMBIA: R. D. Okanagan-

Similkameen, Penticton, Dog Lake, September 23, 1927, coll. W. Mathers (3 g, 1 Y,

OSAC; 1 1 9 TRSC); R. D. Okanagan-Similkameen, Summerland, August 15, 1955,

coll. M. D. Proverbs, feeding on immature Phenacaspispinifolia (4 o, 3 9, CNCI; 1 o, 1

STRSC); R. D. Okanagan-Similkameen, Summerland, August 21, 1964, Pinus

ponderosa (1 Y, CNCI); R. D. East Kootenay, Wardner, August 21, 1977, coll. B. F. & J.

L. Carr (1 Y, CNCI); UNITED STATES: ARIZONA: Apache Co., Adamana, May 7,

1903, coll. H. S. Barber (2 o, USNM); Apache Co., Canyon de Chelly, May 30, 1974,

coll. K. Stephan (1 o, LSAM); Apache Co., Jct. 1-40 & Hwy. 191, August 20-22, 1999,

coll. E. Riley & M. Yoder (1 o, TAMU); Cochise Co., Dragoon Mts. Wood Canyon,

April 29, 1972, coll. K. Stephan (1 Y, FSCA); Cochise Co., Cochise Stronghold

Recreation Area, August 3, 1990, coll. B. F. & J. L. Carr (1 o, CNCI); Coconino Co.,

Fredonia, August 9, 1966, coll. B. F. & J. L. Carr (1 o, 1 9, CNCI); Coconino Co.,

Walnut (1 SEMC); Coconino Co., Walnut, coll. Wickam (1 o, SEMC); Coconino Co.,

Walnut, July, 1922, coll. Wickam (1 o, SEMC); Coconino Co., Williams, May 26, coll.

Barber & Schwarz (1 S, USNM); Gila Co., Roosevelt Reservoir, September 29, 1980,

coll. B. F. & J. L. Carr (1 o, CNCI); Graham Co., S. Graham Mts. 5000ft, July 20, 1974,

coll. K. Stephan (1 o, LSAM; 3 $,1 Y, FSCA); Maricopa Co., Apache Lake, September

23, 1989, coll. B. F. & J. L. Carr (1 o, CNCI); Maricopa Co., Phoenix, January 24, 1930,

coll. S. Flanders (1 o, UCRC); Mohave Co., Hot Springs, June 25, coll. Barber &

Schwarz (2 o, USNM); Navajo Co., Winslow, July 31, coll. Barber & Schwarz (2 g,

USNM); Pima Co., Greaterville, October 8, 1980, coll. B. F. & J. L. Carr (1 g, CNCI);

Pima Co., Vail, May 3, 1975, coll. K. Stephan (1 o, FSCA); Pima Co., Tucson,









December 21, coll. Hubbard & Schwarz (2 g, 3 Y, USNM); Pima, Co., Tucson, January

5, coll. Hubbard & Schwarz, Prosopisjuliflora (1 Y, USNM); Pinal Co., Oracle, January

9, June 29, July 5, 6, coll. Hubbard & Schwarz (3 o, 8 Y, USNM); Pinal Co., Superior,

September 22, 1989, coll. B. F. & J. L. Carr (1 CNCI); Santa Cruz Co., Santa Rita

Mts., 5000 to 8000 ft., July, coll. F. H. Snow (1 SEMC); Santa Cruz Co., Santa Rita

Mts., June 20, coll. Hubbard and Schwarz (1 Y, USNM); Yuma Co., Ft. Yuma, January

21, coll. Hubbard & Schwarz (1 Y, USNM); CALIFORNIA: Butte Co., Chico, May 14,

1942, coll. E. H. Fosen (5 o, 1 9, USNM; 1 o, 1 9, TRSC); Colusa Co., Letts Lake,

April 30, 1980, coll. Fred G. Andrews, S. Kuba, & T. D. Eichlin (1 g, 1 9, CSCA);

Contra Costa Co. Mt. Diablo, April 5, 1952, coll. R. Schuster (3 g, 5 Y, EMEC; 1 g, 1

Y, TRSC); Contra Costa Co., 3mi S. Martinez, November 26, 1987, coll. L. G. Varela, on

pear foliage (27 o, 16 Y, EMEC; 3 o, 4 Y TRSC; 1 o, 3 Y FSCA); Contra Costa Co.,

Albany, reared in Albany insectary (2 o, EMEC); Fresno Co., Selma, May 11, 1950, coll.

P. DeBach, on red scale on grapefruit (1 Y, UCRC); Fresno Co., Fresno, October 16,

1951 (2 o, EMEC); Imperial Co., Brawley, Apr. 13, 1976, ex. Lemon trees (1 g, 1 ,

CSCA); Imperial Co., El Centro, May 4, 1951, coll. B. H. Harrigan, on red scale (1 ,

UCDC); Imperial Co., Brawley, February 26, 1959, coll. E. I. Schlinger, collected by

vacuum insect net in alfalfa field (1 o, UCRC); Imperial Co., Brawley, January 29, 1959,

coll. E. I. Schlinger, Medicago sativa (1 Y, UCRC); Imperial Co., Imperial Valley, June

11, 1971, coll. Flanders, on citrus (1 o, 1 Y, UCRC); Inyo Co., Bishop, July 26, 1921,

coll. L. L. Muchmore, sage (4 o, 3 Y, LACM; 1 o, TRSC); Inyo Co., Westgard Pass,

August 20, 1960, coll. E. I. Schlinger, on Chllvih, 11iuu, (2 g, 2 Y, UCRC; 1 g, TRSC);

Kern Co., 12mi. NW Rosamond, July 22, 1955, coll. R. A. Flock, Lycium cooper A.









Gray (1 Y, UCRC); Kern Co., 7mi. NW Mohave, July 16, 1965, coll. C. W. O'Brien, ex.

Lycium cooper (3 g, 5 9, EMEC; 4 S, 4 9, FSCA; 1 o, 1 9, TRSC); Kern Co., Ft.

Tejon, July 3, August 3, 1930 (9 o, 8 9, TAMU); Los Angeles Co., July, coll. Coquillett

(1 USNM); Placer Co., August, coll. A. Koebele (1 g, USNM); Placer Co., Towle,

Mar. 7, 1913, ex. Aulacaspis manzanitae (1 9, CSCA); Plumas Co., Lake Almanor, Apr.

23, 1971, coll. Fred G. Andrews, ex. pine litter (1 Y, CSCA); Los Angeles Co., Pomona

(1 o, 1 9, LACM; 2 ', INHS); Los Angeles Co., Arroyo Seco, February 21, 1971, coll.

A Mayor (1 o, UCRC); Los Angeles Co., Little Rock, September 3, 1944, April 8, 1945,

coll. G. P. Mackenzie (3 g,UCRC); Los Angeles Co., Los Angeles, February 1944, coll.

R. H. Smith (4 9, UCRC; 1 o, TRSC; 1 9, FSCA); Los Angeles Co., Vincent, August 4,

1952, coll. Timberlake, on Atriplex canescens (1 9, UCRC); Merced Co., Merced, July

18, 1949, coll. R. L. Doutt, ex. Lepidosaphesficus (1 o, EMEC); Monterey Co., UC Big

Creek Reserve, Big Devils Ck. confluence, 36.0770 N 121.594 W, May 26-27, 2002,

coll. S. Lew (1 g, SBMN); Orange Co., Santa Ana, December 18, 1936, coll. C. E.

Norland, Diaspis echinocacti (3 o, 4 9, LACM; 1 o, 1 9, TRSC); Orange Co., Costa

Mesa, January 21, 1943, (rest illegible) (1 o, UCRC); Placer Co., February 7, 1913, coll.

B. B. Whitney (1 o, 2 9, UCRC); Riverside Co., Mecca, February 25, 1914, coll. J. D.

Neils, feeding on Parlatoria blanchardi (1 g, USNM); Riverside Co., Indio, May, 1926,

coll. F. S. Stickney, eating Marlatt scale (6 o, 6 9, USNM); Riverside Co., Mecca, April

17, 1967, coll. B. F. & J. L. Carr (1 3 9, CNCI); Riverside Co., Ripley, June 25, 1946,

coll. W. F. Barr, Pluchea sericea (1 WFBM); Riverside Co., Riverside, March 29,

1989, coll. F. D. Bennett, cactus (4 FSCA); Riverside Co. Oasis, July 1967, coll. Fred

G. Andrews (1 Y, CSCA); Riverside Co., 31mi NBlythe, Apr. 27, 1978, coll. A. R.









Hardy and Fred G. Andrews, collected on Pluchea sericera (1 Y, CSCA); Riverside Co.,

5 mi. N Aguanga, 1160 52' 30" W 33 32' 30" N, August 17, 1978, coll. J. B. Woolley (2

Y, UCRC); Riverside Co., Riverside, August 23, 1965, coll. M. E. Irwin (2 Y, UCRC);

Riverside Co., Mecca, November 17, 1970, coll. R. C. Dickson, on sticky board (2 ', 2

Y, UCRC; 1 o, TRSC; 1 Y, FSCA); Riverside Co., Riverside, August 1, 1924, feeding

on Aleyrodids, (rest is illegible) (2 o, UCRC; 1 o, 1 9, TRSC); Riverside Co.,

Riverside, March 1926, on Aphidiotus on walnut, (rest is illegible) (1 ', 2 9, UCRC);

Riverside Co., Riverside, January 20, 1927, coll. Timberlake (rest illegible) (1 Y,

UCRC); Riverside Co., Coachella Valley, February 23, 2000, coll. G. R. Ballmer, on

lettuce (1 o, UCRC); Riverside Co., Whitewater Canyon, 650m, 33 57' 18" N 1160 38'

39" W, September 4, 1999, coll. Yanega and Gates (1 Y, UCRC); Riverside Co., Indio,

Imi. E. Jefferson Street, Kennedy Ranch, July 1963, coll. H. T. Reynolds, ex. cotton

suction machine (2 o, UCRC); San Bernardino Co. 2mi. W. Phelan, June 7, 1958, coll.

E. I. Schlinger, ex. scale on manzanita (1 Y, UCRC); San Bernardino, Morongo,

September 20, 1944, on Acacia greggi, (rest is illegible) (2 o, UCRC); San Diego Co.,

San Diego, 1922, coll. Armitage, on cactus (2 o, 2 Y, UCRC); San Luis Obispo Co.,

Cuyama River, nr. Cuyama, August 6, 1999, coll. G. R. Ballmer, on Atriplex canescens

root grown (1 Y, UCRC); San Luis Obispo Co., Carrizo Plain N. M., Caliente Ridge,

35012'27" N 1190 82'88" W, December 5, 2003 January 1, January 1-24, February 29-

March 17, March 17-April 2, 2004, coll. M. Caterino, malaise, flight intercept, unbaited

pitfall (2 g, 2 Y, SBMN; 3 o, TRSC); San Luis Obispo Co., Carrizo Plain N. M., Selby

Campground, 3512'27" N 119082'88" W, February 7-29, January 1-24, February 29-

March 17, 2004, coll. M. Caterino, flight intercept (1 o, 2 Y, SBMN); Santa Clara Co.,









San Jose, June (1 o, OSUC); Santa Clara Co. (1 Y, OSUC); Sonoma Co., Healdsburg,

Apr. 21, 1971, ex. Prunus (1 o, CSCA); Tehama Co., Mill Creek, August 31, 1947, coll.

Timberlake, (rest is illegible) (1 Y, UCRC); Tulare Co., Visalia, April 21, 1930, collected

from sugar pine (2 o, LACM); Yuba Co., coll. E. J. Branigan (2 g, 2 Y, UCRC; 1 g, 1

Y, TRSC); COLORADO: Las Animas Co., June 16, 1982, coll. B. F. & J. L. Carr (1 g,

CNCI); IDAHO: Boise Co., Imi. S Ten Mile, June 14, 1988, coll. A. Allen, sweeping (1

o, AAIC); Canyon Co., Parma, 2224 ft., June 15, 1929, coll. C. Wakeland (1 ,

WFBM); Canyon Co., 4.8 NW Walters Ferry, May 16, 1995, coll. W. F. Barr, sweeping

Atriplex canescens & confertifolia (1 Y, WFBM); Elmore Co., Mt. Home, 3138 ft., July

31, 1952, coll. W. F. Barr, Artemisia (1 OSAC); Fremont Co., Warm River Camp

Campground, June 7, 1986, coll. B. F. & J. L. Carr (1 g, CNCI); Latah Co., Moscow,

April 17, 1994, coll. M. M. Fumiss (1 g,WFBM); Owyhee Co., Harpers Fairy Crossing,

August 20, 1979, coll. A. Allen, on sage brush (1 Y, AAIC); Twin Falls Co., Shoshone

Falls, July 10, September 21, 1975, coll. A. Allen, sweeping sagebrush (3 Y, AAIC);

Twin Falls Co., August 5, 1978, coll. A. Allen, sweeping Artemisia (1 Y, AAIC); Twin

Falls Co., Derkies Lake, Snake River Canyon, June 29, 1977, coll. A. Allen, sweeping

sagebrush (1 o, AAIC); MONTANA: Roosevelt Co., Snowden Bridge, June 11, 1991,

coll. D. L. Gustafson (1 o, MTEC); Powder River Co., 5mi W. Broadus, June 7, 1999,

coll. D. L. Gustafson (1 o, MTEC); Madison Co., 5mi E. Norris, Bear Trap Prim. Area

Madison, 5000 ft., July 26, 1986, S. M. Fondriest (1 Y, MTEC); NEVADA: Elco Co.,

Ruby Mtns. 7000 ft., May 14, 1975, coll. James H. Baker (4 g, USNM); NEW

MEXICO: (1 g, 1 Y, INHS); Eddy Co., Lincoln National Forest, 4.5mi SW Queen,

Hwy. 137, 1675m, 3212'01" N 104 40'10" W, August 15-25, 2001, coll. J. C. Schaffner









(1 TAMU); Socorro Co., Beartrap Canyon, Mt. Withington, July 31, 1990, coll. B. F.

& J. L. Carr (1 Y, CNCI); OKLAHOMA: Latimer Co., March 1986, coll. K. Stephan (2

o, TAMU); OREGON: Klamath Co., Klamath Falls, June 6, 1956, coll. Joe Schuh (1 g,

3 Y, OSAC); Klamath Co., Upper Klamath Lk., Geary Canal, May 23, 1958, coll. Joe

Schuh (1 o, OSAC); Wasco Co., Bear Springs, October 6, 1940, coll. K. M. & I. M.

Fender (1 o, OSAC); TEXAS: Brewster Co., Big Bend Nat. Pk., 14mi E. Pantherjct.,

September 9, 1985, coll. W. F. Barr, Croton (1 o, WFBM); Cameron Co., Brownsville,

January 18, 1915, coll. Timberlake, assoc., with Dactylopius (1 o, 1 Y, UCRC); Gaines

Co., Seminole, June 13, 1983, coll. B. F. & J. L. Carr (1 Y, CNCI); Hudspeth Co., Indio

Mountains Research Station, vic. Indio ranch house, 4040 ft., 30o46'37" N 10500'43"

W, June 12-13, 2002, coll. E. G. Riley, R. Diaz & M. J. Yoder (1 g, 2 Y, TAMU; 1 g, 1

Y, TRSC); Hudspeth Co., Indio Mountains Research Station, Cougar Canyon, 4040 ft.,

30o46'59" N 105001'06" W, March 30-April 12, 2002, coll. A. R. Gillogly, Malaise Trap

(1 TAMU); Hudspeth Co., Indio Mountains Research Station, Squaw Spring,

30o47'49" N 10500'43" W, April 12-13, 2002, coll. E. G. Riley, & M. J. Yoder (1 ,

TAMU); Hudspeth Co., Indio Mountains Research Station, Squaw Creek, 4200 ft.,

30o47'49" N 10500'43" W, March 30-April 12, 2002, coll. R. Caesar & A. R. Gillogly

(1 2 9, TAMU); Kennedy Co., 13.5mi S Sarita, October 11, 1994, coll. E. G. Riley (1

Y, TAMU); Pecos Co., Hwy. 385 rest stop, 28mi S Ft. Stockton, April 19, 1997, coll. E.

Riley (1 o, TAMU); Presidio Co., Big Bend Ranch St. Nat. Ar., Aqua Adento, June 18-

23, 1990, coll. D. Judd, Malaise Trap (1 o, 7 Y, TAMU; 1 o, 1 9, TRSC); San

Augustine Co., 10mi SE Broaddus, Coleman Cemetery, March 19, 1994, coll. E. Riley (1

Y, TAMU); Travis Co., vic. Long Hollow Ck., 30o27'43"9752' 19", May 8, 26, 1993,









coll. Alexander, Quinn, Riley, Wharton, et al., on Juniperus ashei (3 o, 4 9, TAMU; 1

o, 1 9, TRSC); Travis Co., vie. Long Hollow Creek and Cypress Creek,

30o27'43"97o52'19", 30025'58"97052'01", March 26, June 18-19, 1994, coll. M. Quinn,

E. Riley, R.Wharton, on Quercus buckleyi (2 Y, TAMU); Val Verde Co., Amistad

Reservoir, Hwy 406, June 2, 2000, coll. E. G. Riley (3 o, 4 9, TAMU; 1 o, TRSC);

UTAH: Washington Co., St. George, coll. Wickham (1 g, SEMC); WASHINGTON:

Benton Co., Hanford Works, 640 ft., July 30, 1952, September 17, 1952, September 24,

1952, coll. R. H. Whittaker, Sagebrush (3 Y, OSAC).

Remarks. Specimens are most often collected while sweeping brush, especially

Artemisia spp. This beetle seems to occupy a similar niche to that of C. randalli, which

is often collected in the same areas and on the same plants. While the distribution of this

beetle is quite extensive, it does not seem to occur east of the Mississippi River.

Cybocephalus kathrynae T. R. Smith, New Species

(Figs. 4-11, 4-12-4-16)

Diagnosis. Male and female black. Antennal club has a smooth margin without a

serrated edge and the scutellum has concave margins (Figs. 4-11), distinguishing this

species from C. randalli, C. californicus, C. nigritulus and C. nipponicus.

Etymology. This species is named for Kathryn Lang Zara-Smith, wife of the

species name's author.

Description. Male. Form: Elongate oval; strongly convex dorsally. Length: 1.3-

1.6 mm (measured from apex of clypeus to apex of elytra); width: 0.8-1 mm (measured at

base of elytra). Color: Head, thorax, elytra and underside dark brown or black, the

extreme edge of elytral apices with brown border. Front legs and antennae light brown or

brown, middle and hind legs black or dark brown. Head: Broad and convex, clypeus









wide and moderately produced. Eyes large and tear-shaped with internal margins

distinct. Genae just visible from above and slightly explanate. Dorsal surface smooth but

distinctly alutaceous with uniform punctation. Antennae 11-segmented including a club

with 3 antennomeres, club length about 12-2/3 the size of eye. Club flat and distinctly

separated from funicle and with a smooth margin. First club antennomere wider than

long, second club antennomere larger than either first or third club antennomere and

about as long as wide. Terminal club antennomere emarginate (Fig. 4-12). Antennomere

3 as long or longer than 4 and 5 combined. Pronotum: Strongly convex and shiny,

lateral margin curved; anterior angle more narrowly arcuate than posterior. Surface

evenly punctured and alutaceous. Scutellum: Alutacous and triangular with concave

margins. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex, sides

almost parallel but slightly sinuous, apices rounded, length shorter than combined width

(29:38). Very alutaceous along dorsal surface and distinctly punctured, punctation

ending before base of elytra, forming a narrow impunctate area at base and sides.

Median margin and edge of elytra bordered. Distinct striations at apices of elytra.

Underside: Metasternum extremely alutaceous and roughly punctate, and clothed in long

coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs thinly

covering the surface. Legs: All femora glossy with short hairs. Profemora narrowing

slightly at basal end, meso- and metafemora extremely flattened and expanded. All tibiae

slightly but distinctly curved and dilated toward the apex. Protibiae with short hairs

along outer margin. Meso- and metatibiae with long stiff hairs along outer margin. Four

tarsomeres, claw tarsomere as long or almost as long as 2 tarsomeres preceding it.

Median lobe: Sides curving and convergent forming a sharp point (Fig. 4-13). In profile,









curved (Fig. 4-14). Median plate on surface very slightly elevated. Basal plate: Sides

convergent and emarginate at apex (Fig. 4-15).

Female. Nearly identical to male.

Geographic distribution. Known only from the southern coast of Florida (Fig. 4-

16).

Hosts. Cybocephalus kathrynae has been collected in very close proximity to

Haliaspis nakaharai Howell and Haliaspis uniolae Takagi (Diaspididae). While these

beetles have not actually been observed feeding on the aforementioned scale species, they

were the only scale species in the beetle's habitat and exhibited signs of predation.

Type material examined. The holotype, deposited in the MCZC, is a male

specimen glued to a point with the following labels: USA: Florida, Monroe Co., Bahia

Honda State Park southeast end of island N2439'54"-W81015'21" (printed) [white

rectangular label] / V-14-2005, Trevor Smith & R. D. Cave, sifting sand dune leaf litter

around Uniolapaniculata (printed) [white rectangular label] / HOLOTYPE

Cybocephalus kathrynae T. R. Smith Det: Trevor Smith (printed) [red rectangular label].

The designated allotype, deposited in the MCZC, is a female specimen glued to a point

with the following labels: USA: FL: Monroe Co. Bahia Honda St. Pk. SE. end of island

3-III-2005, Trevor Smith, M. C. Thomas, sand litter, primary dune (printed) [white

rectangular label] / ALLOTYPE Cybocephalus kathrynae T. R. Smith Det: Trevor Smith

(printed) [blue rectangular label]. Designated paratypes are as follows: UNITED

STATES: FLORIDA: Monroe Co., Bahia Honda State Park, southeast end of island,

December 1, 1999, coll: Paul Skelley, berlese litter under trees of 20 dunes (1 g, TRSC);

Monroe Co., Bahia Honda State Park, southeast end of island, N2439'54"-W81015'21",









May 14, 2005, coll: Trevor Smith & R. D. Cave, sifting sand dune leaf litter around

Uniolapaniculata (3 ', 4 9, FSCA); Miami-Dade Co., Key Biscayne, Bill Baggs Cape

Florida State Park., N25040'08"-W80009'17", June 30, 2005, coll: Trevor Smith, sifting

sand dune leaf litter (1 o, 4 Y, FSCA).

Remarks. This species was collected by sifting sand and leaf litter in primary and

secondary sand dunes where sea oats (Uniolapaniculata L.) and seashore dropseed

(Sporobolus virginicus (L.)) were present. The sea oats were infested with H. uniolae

and the seashore dropseed was infested with H. nakaharai. It has only been collected on

Bahia Honda Key in Monroe County and in Bill Baggs Cape Florida State Park in

Miami-Dade County but may occur in other areas of Florida with naturally occurring

sand dunes where sea oats are present and infested with armored scales.

Cybocephalus nigritulus LeConte

(Figs. 4-17-4-22)

Cybocephalus nigritulus LeConte 1863: 64.

Diagnosis. Male and female black and very glossy. Size similar to C. californicus

but slightly larger. Antennal club smaller than eye and truncate at terminal antennomere;

each antennomere of club distinctly separated forming a serrated edge, distinguishing this

species from C. kathrynae. The truncate terminal club antennomere distinguishes this

species from C. randalli which has a rounded terminal antennomere. Protibia more

dilated than in C. californicus and the translucent impunctate area at the apices of elytra

much smaller than in C. californicus. Male basal plate and median lobe are unique and

easily distinguished from all other species.

Redescription. Male. Form: Elongate, oval; contractile; strongly convex

dorsally. Length: 1.0-1.55 mm (measured from apex of clypeus to apex of elytra);









width: 0.85-1.19 mm (measured at base of elytra). Color: Head, thorax, elytra glossy

black, underside black, with lateral margin of pronotum and apical margin of elytra

yellowish and translucent. Front legs and antennae amber, middle legs amber or light

brown, hind legs brown or dark brown. Head: Broad and convex, clypeus moderately

produced. Eyes tear-drop shaped and large with internal margins distinct. Genae not

visible from above but slightly explanate when viewed laterally. Dorsal surface smooth,

but under high magnification very finely alutaceous, finely punctured. Antennae 11-

segmented including a club with 3 antennomeres; club length, 12-2/3 the width of eye; club

flat and distinctly separated from funicle and with a distinctly serrated margin. First club

antennomere wider than long, second club antennomere much larger than first and third

club antennomeres and about as long as wide; terminal antennomere of club truncate,

setose, and about as long as wide (Fig. 4-17). Antennomere 3 about the same length as

antennomeres 4 and 5 combined. Pronotum: Strongly convex, under high magnification

finely alutaceous with extreme lateral edges curving slightly and almost straight, anterior

angle more narrowly arcuate than posterior. Surface punctation uniform, but distinct, and

somewhat sparse, surface shiny. Scutellum: Alutaceous and triangular with slightly

convex margins. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex,

sides almost parallel but slightly sinuous, apices rounded. Length somewhat shorter than

combined width (34:44). Punctation distinct except along extreme edge of apex which is

smooth and without punctation; under high magnification finely alutaceous. Median

margin and apex of elytra bordered. Underside: Metasternum extremely alutaceous,

roughly punctured and clothed in long coarse hairs. Abdominal sternites alutaceous and

punctate with long coarse hairs thinly covering surface. Legs: Femora glossy, broad,









flattened and sparsely covered with short hairs. Pro- and mesofemora about same width

from end to end. Metafemora expanded in middle. Tibiae slightly but distinctly curved

and dilated towards apex. Protibiae much more dilated than others and with short hairs

along outer margin. Meso- and metatibiae with long stiff hairs along outer margin. Four

tarsomeres, claw tarsomere as long or almost as long as 2 preceding tarsomeres

combined. Median lobe: Sides parallel or slightly convergent curving into a large

triangular point (Fig. 4-18, 4-19). In profile strongly curved from middle (Fig. 4-20).

Median plate on surface elevated. Basal plate: Sides parallel sides at base, very slightly

tapering towards apex. Apex flat with a distinct triangular protuberance in center (Fig. 4-

21).

Female. Nearly identical to male.

Geographic distribution. Alabama, Florida, Georgia, Indiana, Louisiana,

Massachusetts, Michigan, Minnesota, Mississippi, Ontario, Pennsylvania, Rhode Island,

South Carolina (Fig. 4-22).

Hosts. Like all cybocephalids, the primary food source of these beetles is armored

scale insects (Diaspididae). Cybocephalus nigritulus has been seen in association with or

reported feeding on C. pinifolia (Riley 1882), Fiorinia theae Green (Flanders 1934; from

label data), Pseudaulacaspis cockerelli (Cooley) (from label data), and Pseudaulacaspis

pentagon (Targioni-Tozzetti) (Collins and Whitcomb 1975).

Type material examined. The holotype in the MCZC is a male specimen glued to

a point with the following labels: orange disc / Type (printed) 6987 (handwritten) [dark

orange rectangular label] / Cybocephalus nigritulus Lec. (handwritten) [white rectangular

label] / Cybocephalus nigritulus LeConte Det: Trevor Smith (printed) [white rectangular









label]. The orange disc indicates that the specimen was collected in the southern or gulf

states. Two specimens in the MCZC that are possibly syntypes were also examined. One

male and one female are glued to points on separate pins, but have the same labels as

follows: Detroit, Mich. (printed) [white rectangular label] / Cybocephalus nigritulus

LeConte Det: Trevor Smith [white rectangular label]. These specimens may be the

Michigan specimens referred to by Horn (1879) in his redescription of this species.

Other material examined. CANADA: ONTARIO: Kent Co., Tilbury, August

1967, coll. K. Stephan, sifting (1 Y, FSCA); Essex Co., Wheatley, June 1967, coll. K.

Stephan, coccids on aspen (1 6, FSCA); UNITED STATES: ALABAMA: Walker Co.,

Jasper, September 23, 1979, coll. Tim King, at light (1 o, AAIC); FLORIDA: Alachua

Co., Gainesville, San Felasco Hammock St. Pres., Apr. 10, June 10, June 26, 2004, coll.

Trevor Smith, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (20 6, 20

9, TRSC; 8 6, 10 9, FSCA); Alachua Co., Gainesville, University of Florida Natural

Area., March 12, 2005, coll. Trevor Smith, feeding on Pseudaulacaspis cockerelli on

Magnolia grandiflora (20 o, 20 9, FSCA); Alachua Co., Gainesville, Haile Plantation,

March 30, 2005, coll. Trevor Smith, feeding on Pseudaulacaspis cockerelli on Magnolia

grandiflora (5 o, 5 9, FSCA); Alachua Co., Gainesville, Wilmont Gardens, March 8,

1974, coll. Fred Collins, on Camellia (1 Y, USNM; 1 Y, FSCA); Alachua Co.,

Gainesville, July 18, 1964, coll. R. E. White (1 Y, USNM); Alachua Co., Gainesville,

July 18,19, May 23, 24, 1964, coll. R. E.White (5 6, FSCA); Alachua Co., Gainesville,

Devil's Millhopper, September 28, 1997, coll. Vince Golia, beating trees (1 Y, FSCA);

Alachua Co., Gainesville, May 28, 1988, coll. F. D. Bennett, pred. on Fiorina theae on

Camellia (1 6, FSCA); Alachua Co., N. E. Gainesville, May 16, 2000, coll. J. Eric









Cronin, feeding on Pseudaulacaspis cockerelli on Magnolia grandiflora (1 g, 1 9,

FSCA); Alachua Co., Gainesville, San Felasco Hammock St. Preserve, March 20, 2004,

coll. M. C. Thomas, ex. Magnolia sp. (1 Y, FSCA); Brevard Co., Titusville, SR 405,

Enchanted Forest Sanct., White Trail, September 14-28, 2000, January 15-31, 2001, coll.

Z. Prusak, P. J. Russell, S. M. Fullerton, malaise trap, xeric oak hammock (1 g, 1 9,

UCFC); Dade Co., Miami, Snapper Creek Plaza, October 20, 1991, coll. F. D. Bennett,

Pseudaulacaspis cockerelli / Strelitzia ( 1 FSCA); Gadsen Co. Quincy, Hyw. 267,

March 6, 1974, coll. Fred Collins, on Camellia (1 g, USNM); Gadsen Co., March 6,

1974, sawdust, Melia azedarach (1 Y, FSCA); Orange Co., Orlando, UCF, April 5, 1999,

coll. P. Russell, S. Fullerton, malaise trap, pond pine community/dahoon holly (2 9,

UCFC); Orange Co., Orlando, UCF, April 19, 1999, coll. P. Russell, S. Fullerton, malaise

trap, maidencane marsh (2 Y, UCFC); Orange Co., Orlando, UCF, May 11, 17, 24, June

2, 1999, coll. P. Russell, S. Fullerton, malaise trap, cypress forest (1 g, 3 9, UCFC);

Orange Co., Orlando, UCF, July 2, 1997, coll. S. Fullerton, malaise trap, long leaf

pine/sand pine/turkey oak (1 o, UCFC); GEORGIA: Berrien Co., Alapaha, May 6,

2004, coll. Trevor Smith, on Magnolia grandiflora feeding on Pseudaulacaspis

cockerelli (2 o, 1 9, TRSC); LOUISIANA: East Baton Rouge Parish, Baton Rouge,

Bluebonnet Swamp, August 18, 2000, coll. A. R. Cline, my light (1 Y, LSAM);

MASSACHUSETTS: Suffolk Co., Boston, Arnold Arboretum, July 14, 1921, coll.

Harold Morrison, swept from 5-leaf pines behind lab (1 g, USNM); MISSISSIPPI:

Smith Co., July 22, 1956 (1 9 TRSC); PENNSYLVANIA: Allegheny Co. (1 g, SEMC);

SOUTH CAROLINA: Dorchester Co., Summerville, April 2, 1909, coll. J. G. Sanders,

on tea scales (1 g, USNM).









Remarks. These beetles are usually collected by hand, either beating or simply by

gleaning and aspirating. They are also occasionally picked up in Malaise traps. While

they are sometimes found in urban areas, they seem to be collected more often in forested

areas. The senior author collected hundreds of these beetles by spotting and aspirating

them from Magnolia grandiflora L. infested with P. cockerelli. While this beetle seems

to be very widespread, ranging from Canada to Florida and west to the Mississippi River,

there are relatively few specimens in collections. There is also relatively less known

about the hosts of this particular species.

Cybocephalus nipponicus Endrody-Younga

(Figs. 4-10, 4-23-4-27, 4-29)

Cybocephalus nipponicus Endrody-Younga 197 la: 244-245.

Diagnosis. Male bicolored with yellow or tan head, pro- and mesosternum,

antennae and legs and remainder of the body black, thus distinguishing this species from

all other Cybocephalus in North America. The female is almost completely black with

yellow front legs and antennae, with remaining legs either light brown or brown. The

antennal club is smaller than the eye, truncate and each antennomere is distinctly

separated to form a serrated edge. The male basal plate and median lobe are distinctly

different from other species.

Redescription. Male. Form: Elongate, oval; contractile; strongly convex

dorsally. Length: 1.0-1.35 mm (measured from apex of clypeus to apex of elytra);

width: 0.9-1.30 mm (measured at base of elytra). Color: Head, prothorax, mesothorax,

antennae and legs yellow or yellowish brown; metathorax, elytra and abdomen black.

Head: Broad and convex, clypeus moderately produced and comparatively narrow. Eyes

large, oblong, with internal margins distinct. Genae not visible from above and not









explanate. Dorsal surface smooth, but under high magnification finely alutaceous,

punctation fine. Antennae 11-segmented including a club with 3 antennomeres; club

length short, about 12-2/3 the length of eye. Club flat and distinctly separated from funicle

and with a distinctly serrated margin. All club antennomeres wider than long and

creating a serrated edge on inside margin of club. First club antennomere wider than

long, second club antennomere larger than first and third club antennomeres and slightly

wider than long. Terminal club antennomere truncate, setose, and slightly wider than

long (Fig. 4-23). Third antennomere slightly shorter than segments 4 and 5 combined.

Pronotum: Strongly convex, lateral margins slightly curved or nearly straight; anterior

angle more narrowly arcuate than posterior. Dorsal surface under high magnification

finely alutaceous with fine punctation. Scutellum: Alutaceous and triangular with slightly

convex margins. Elytra: Uniform width narrowing at the apical 1/5. Strongly convex,

sides almost parallel but slightly sinuous, apices rounded. Length somewhat shorter than

combined width (33:40). Fine punctation at the base disappearing completely at distal

edge but toward middle clearly three-armed and finely alutaceous. Median margin and

apex of elytra bordered. Underside: Metasternum extremely alutaceous, roughly

punctured and clothed in long coarse hairs. Abdominal sternites alutaceous and punctate

with long coarse hairs thinly covering surface. Legs: Femora glossy, broad, flattened

and sparsely covered with short hairs. Pro- and mesofemora about same width from end

to end. Metafemora expanded in middle. Tibiae slightly but distinctly curved and dilated

towards apex. Protibiae much more dilated than others and with short hairs along outer

margin. Meso- and metatibiae with long stiff hairs along outer margin. Four tarsomeres,

claw tarsomere as long or almost as long as 2 preceding tarsomeres combined. Median









lobe: Sides parallel at base, extending to a large apical triangle (Fig. 4-24), in profile

strongly curved at tip (Fig. 4-25). Basal plate: Sides parallel at base, tapering apically

(Fig. 4-26).

Female. Similar to male except all black, lateral margin of the prothorax yellow or

translucent and apical margin of elytra and middle and hind legs light brown or brown.

Geographic distribution. Connecticut, Delaware, Florida, Hawaii, Maryland,

Massachusetts, New Jersey, New York, Pennsylvania, Rhode Island, Texas, Virginia,

Washington D.C. (Fig. 4-27), Asia, Europe, Micronesia, West Indies and South Africa.

This is a recently introduced species native to Southeast Asia, therefore specimens are

rarely found in North American collections. For more information on the distribution of

these beetles, see Drea and Carlson (1988), Jefferson et al. 1995, Van Driesche et al.

(1998), Hudson et al. (2000), Deepak et al. (2003), Spichiger (2004), and HDA (2004).

Hosts. This predator has been reported in association with or feeding on a large

number of armored scale insects (Diaspididae): Aonidiella sp. (Endr6dy-Younga 1971a),

Aspidiotus destructor Signoret (Endr6dy-Younga 1971a,b; from label data), Aulacaspis

crawii (Cockerell) (from label data), Aulacaspis yasumatsui Takagi (Heu and Chun 2000;

from label data), Chrysomphalus bifasciculotus Ferris (Hayashi 1978; Tanaka and Inoue

1980), Fiorinia externa Ferris (Spichiger 2004; from label data), Hemichionaspis sp.

(Endr6dy-Younga 1971a), P. cockerelli (Cooley) (from label data), P. pentagon

(Targioni-Tozzetti) (Yasuda 1981; Endr6dy-Younga 1971a,b), Diaspidiotus

macroporanus (Takagi) (Tachikawa 1974), Quadraspidiotus perniciosus (Comstock)

(Alvarez and Van Driesche 1998a), Unaspis euonymi (Comstock) (Alvarez and Van

Driesche 1998b; Drea and Carlson 1988), and Unaspisyanonensis Kuwana (Tanaka and









Inoue 1980). It has also been reported feeding on the citrus red mite, P. citri (Tanaka and

Inoue 1980).

Material examined. UNITED STATES: CONNECTICUT: Hartford Co.,

Windsor, Connecticut Agricultural Experiment Station, Valley Laboratory, Sept. 15,

2005, coll. R. S. Cowles, feeding on Fiorinia externa Ferris on Abiesfraseri (Pursh.) (15

o, 20 9, FSCA); FLORIDA: Collier Co., Naples, Apr. 25, 2002, coll. S. Krueger, on

Cycas sp. (1 o, 3 9, FSCA); Dade Co., Turnpike, Snapper Creek Plaza, June 25, 1990,

coll. F. D. Bennett, ex. Pseudaulacaspis cockerelli on Ravenela madagascarensis (4 o, 2

Y, FSCA); Dade Co., S. Miami, Mar. 24, 2004, coll. P. Duetting, feeding on Aulacaspis

yasumatsui on Cycas revoluta (10 o, 10 9, TRSC); Dade Co., S. Miami, jct. US-1 & SW

141 St., March 24, Apr. 12, 2004 coll. Trevor Smith, feeding on Aulacaspis yasumatsui

on Cycas revoluta (10 o, 10 9, TRSC; 10 o, 10 9, FSCA); Hillsborough Co., Plant City,

May 14, 2004, coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta

(10 g, 10 9, TRSC; 10 g, 10 9, FSCA); Hillsborough Co., Tampa, Downtown, July 14,

2005, coll. Trevor Smith, feeding on Aulacaspis yasumatsui on Cycas revoluta (10 o, 10

9, FSCA); Manatee Co., Terra Ceia, Feb. 3, 1994, coll. M. Runnals, oleander (2 S, 4 9,

FSCA); Orange Co., Orlando, UCF, April 26, 1999, coll. P. Russell, S. Fullerton, malaise

trap, maidencane marsh (1, UCFC); Sarasota Co., Sarasota, 1315 38th St., Jan. 8, 2005,

coll. J. H. Frank, feeding on Aulacaspis yasumatsui on Cycas rumphii (3 o, 3 9, TRSC);

Seminole Co., Longwood, Oct. 7, 2004, coll. Trevor Smith, feeding on Aulacaspis

yasumatsui on Cycas revoluta (7 o, 5 9, TRSC); MARYLAND: Prince George's Co.,

Beltsville, August 4, 1989, coll. R. Hendrickson, J. Drea, ex: Unaspis euonymi on:

Euonymus sp. (3 o, 2 9, TAMU; 1 o, 1 9 TRSC); TEXAS: Brazos Co., Texas A&M









West Campus, September 27, 2000, coll. M. Yoder (1 g, TAMU); INSECTARIES:

CALIFORNIA: Orange Co., Anaheim, January 30, 1935, Insectary (3 g, 3 9, LACM; 1

o, TRSC); Orange Co., Anaheim, October 15, 1952, coll. P. DeBach, on purple scale (1

o, 5 9, UCRC); Orange Co., Santa Ana, July 1, 1944, (rest is illegible) (2 g, 3 9,

UCRC; 1 o, 1 9 TRSC); Orange Co., Insectary, January, 1948 (1 g, UCRC);Ventura

Co., Ventura, Rincon-Vitova Insectaries, Sept. 8, 2004 (10 o, 10 9, TRSC; 10 g, 10 9,

FSCA); NEW YORK: Cayuga Co., Locke, IPM Laboratories Inc., June 2001, ex. China

(1 g, 6 9, FSCA; 1 g, 1 9, LSAM); PENNSYLVANNIA: Delaware Co., Swarthmore,

July 16-20, 1988, coll. Mike Rose, Texas A&M Quarantine Lab (native to Korea) (4 g, 8

9, TAMU);

Remarks. Cybocephalus nipponicus is the only U.S. cybocephalid which has a

sexually dimorphic color pattern (Fig. 4-28). This species, now widely distributed

throughout the eastern U.S., was misidentified as C. binotatus and released in south

Florida in 1998 to combat A. yasumatsui (Anon. 1998; Howard and Weissling 1999) on

sago palms (Cycas spp.). These two species have been confused in the past as evidenced

by Endrody-Younga mixing specimens of C. binotatus (Fig. 4-28) and C. nipponicus

(Fig. 4-29) in the description of C. binotatus in his 1968 monograph of the Palearctic

cybocephalids. He later clarified this in his description of C. nipponicus and a

redescription of C. binotatus (Endrody-Younga 1971a). A male topotype of C. binotatus

was compared to the beetles released in Florida. Two black spots on the yellow

pronotum, a metallic sheen on the elytra, and distinctive male genitalia clearly separate C.

binotatus from C. nipponicus. It was believed that the 1998 release of C. nipponicus

from Thailand was the first introduction of these beetles into the state of Florida (Howard









et al. 1999). However, specimens dating back to 1989 were found in the FSCA and

identified as C. nipponicus.

Notes about type material of C. binotatus. The studied topotype specimen in the

BMNH is a male glued to a card with the following labels: W. Almora, Kumaon, H.G.C.

(printed) [white rectangular label] / Cybocephalus binotatus, Grouv. (printed) [white

rectangular label] / Ent. Mo. Mag. 1923 Det. G.C.C. (printed) [upside down white

rectangular label] / H. G. Champion Coll. B. M. 1953-156 (printed) [white rectangular

label] / Kirejtshuk det 1996 (printed) Cybocephalus binotatus (handwritten) [white

rectangular label]. This specimen is one of six collected by Champion (1923), of which

one specimen of this series was designated as the neotype by Endrody-Younga (1971a)

when he could not find the original holotype deposited in the Assam Coll. (Grouvelle

1908).

Cybocephalus randalli T. R. Smith, New Species

(Figs.4-30-4-36)

Diagnosis. Male and female black. Clypeus much broader and more extended than

in C. californicus. Antennal club is unlike all other North American species in that it is

as large or larger than the eye, and the terminal antennomere of the club is rounded

apically rather than truncate as in C. californicus, C. nigritulus, and C. nipponicus. Each

club antennomere is distinctly separated and forming a serrated edge, unlike C.

kathrynae. Front tibia is quite dilated, much more so than in C. californicus. In males,

the basal plate and median lobe are easily recognizable.

Etymology. This species is named in honor of Randall Edwards Smith, father of

the species name's author.









Description. Male. Form: Elongate oval; contractile; strongly convex dorsally.

Length: 1.2-1.3 mm (measured from apex of the clypeus to apex of elytra); width: 0.85-

0.95 mm (measured at base of elytra). Color: Head, thorax, elytra and underside black,

lateral margin of pronotum and apical margin of elytra yellowish and translucent; legs

and antennae dark brown or black. Head: Broad and convex, clypeus wide and

produced. Eyes comparatively small, oblong with internal margins distinct. Genae not

quite visible from above and slightly explanate. Dorsal surface smooth, under high

magnification distinctly alutaceous with fine punctation. Antennae 11-segmented

including a club with 3 antennomeres; club large and about same size as or larger than

eye. Club flat and distinctly separate from funicle and with a distinctly serrated margin.

First and second club antennomeres wider than long, terminal club antennomere longer

than wide, apically rounded and quite setose (Fig. 4-30). Third antennomere shorter than

antennomeres 4 and 5 combined. Pronotum: Strongly convex and glossy; anterior angle

more narrowly arcuate than posterior. Surface finely punctate and alutaceous.

Scutellum: Triangular with straight or slightly convex margins. Elytra: Uniform width

narrowing at the apical 1/5. Strongly convex, sides almost parallel, apices rounded, length

somewhat shorter than combined width (30:38). Very sparsely punctate and distinctly

alutaceous, punctation ending before the base of elytra, smooth at sides and base.

Median margin and apex of elytra bordered. Apices of elytra with distinct striations (Fig.

4-35). Underside: Metasternum extremely alutaceous and roughly punctate, and clothed

in long coarse hairs. Abdominal sternites alutaceous and punctate with long coarse hairs

thinly covering the surface. Legs: All femora glossy, broad, flattened and sparsely

covered with short hairs. Pro- and mesofemora about the same width from end to end.









All tibiae slightly but distinctly curved and dialated toward the apex. Metafemora

expanded in the middle. Protibiae much more dilated than the others and with short hairs

along the outer margin. Meso- and metatibiae with long stiff hairs along the outer

margin. Four tarsomeres, claw tarsomere as long or almost as long as 2 preceding

tarsomeres combined. Median lobe: Sides parallel before curving into a triangular apex

(Fig. 4-31, 4-32). In profile, slightly curved at tip (Fig. 4-33). Median plate not elevated.

Basal plate: Sides parallel at base, narrowing slightly before rounding at apex with a

large concavity in the center (Fig. 4-34).

Female. Nearly identical to male.

Geographic distribution. California, Idaho, Nevada, Utah, Washington (Fig. 4-

36).

Hosts. Specimens of C. randalli were collected while sweeping Artemisia spp.,

therefore it is a distinct possibility that these beetles feed on a scale found on Artemisia.

Type material examined. The holotype, deposited in the MCZC, is a male

specimen glued to a point with the following labels: USA: IDAHO FALLS CO.

SHOSHONE FALLS 14 VII 1975 LEG: A ALLEN SWEEPING SAGE BRUSH

(printed) [white rectangular label] / HOLOTYPE Cybocephalus randalli T. R. Smith Det:

Trevor Smith (printed) [red rectangular label]. The designated allotype, deposited in the

MCZC, is a female specimen glued to a point with the following labels: Calif: Inyo Co.

Saline Valley Dunes III-30-1976 cereal bowl pit trap D. Giuliani (printed) [white

rectangular label] / ALLOTYPE Cybocephalus randalli T. R. Smith Det: Trevor Smith

(printed) [blue rectangular label]. Designated paratypes are as follows: UNITED

STATES: CALIFORNIA: San Benito Co., 8.2 mi on Panoche rd. from 1-5, Feb. 4, 1979









- Mar. 15, 1979, coll. A. J. Gilbert, Ethylene glycol pit trap in Ephedra area (1 o, TRSC);

San Benito Co., 8.2 mi on Panoche rd. from 1-5, March 15 April 14, 1979, coll. A. J.

Gilbert, antifreeze pit trap (1 Y, CSCA); Riverside Co., Slover Ave. dump, 315m, 34 03'

56" N 1170 21' 32" W, April 24, 2001, coll. Hawks and Yanega (1 g, UCRC); IDAHO:

Twin Falls Co., Shoshone Falls, July 10, 1975, coll. A. D. Allen (1 o, USNM); Twin

Falls Co., Shoshone Falls, September 21, 1975, coll. A. Allen, sweeping sagebrush (1 o,

TRSC; 1 Y, AAIC); NEVADA: Churchhill Co., 6mi East of Frenchman, August 22,

1972, coll. Stephen J. Chaplin (1 Y, USNM; 1 Y, TRSC); Nye Co., Mercury, July 21,

1964, onAritri (1 o, USNM); Nye Co., Mercury, February 7, 1961 (1 9, USNM);

UTAH: Washington Co., St. George, July, coll. Wickham (1 Y, CNCI); Washington Co.,

St. George (1 Y, MTEC); WASHINGTON: Grant Co., Smyrna, June 19, 1932, coll. M.

H. Hatch (1 o, FSCA);

Remarks. This species is less frequently collected than the sympatric C.

californicus. Whether this beetle is actually less common than C. californicus or whether

they just occupy a slightly different niche is unknown. While these beetles are usually

picked up by sweeping, they occasionally show up in pitfall traps. This is probably a

case of the optimal collecting technique not being employed to obtain this species.




























Figure 4-1. Complete Cybocephalus nipponicus genitalia: ED = ejaculatory duct; IS =
internal sac; MA = muscle attachment; ML = median lobe; MS = median
strut; Tbp = tegmen basal plate; Tdp = tegmen dorsal piece; TS = tegminal
strut.


Figure 4-2. Lateral habitus of Cybocephalus randalli.




















4-3 4-4 4-5













4-6 4-7 4-8

Figure 4-3. Truncate antenna of C. californicus.

Figure 4-4. Emarginate antenna of C. californicus.

Figure 4-5. Median lobe, dorsal view, of C. californicus.

Figure 4-6. Median lobe, dorsal view (slide mounted), of C. californicus.

Figure 4-7. Median lobe, lateral view, of C. californicus.

Figure 4-8. Basal plate, ventral view, of C. californicus.










































Figure 4- 9. U.S. states and Canadian provinces from which specimens of Cybocephalus
californicus have been collected.

















































Figure 4-10. Scutellum of Cybocephaus nipponicus.

Figure 4-11. Scutellum of Cybocephalus kathrynae.



















4-12


4-13


4-14 4-15

Figure 4-12. Antenna of C. kathrynae.
Figure 4-13. Median lobe, dorsal view (same as slide mounted), of C. kathrynae.
Figure 4-14. Median lobe, lateral view, of C. kathrynae.
Figure 4-15. Basal plate, ventral view, of C. kathrynae.




























) Key
Biscayne


Honda Key


Figure 4-16. Collection localities of Cybocephalus kathrynae in Florida.




















4-17 4-18













4-19 4-20 4-21

Figure 4-17. Antenna of C. nigritulus.

Figure 4-18. Median lobe, dorsal view, of C. nigritulus.

Figure 4-19. Median lobe, dorsal view (slide mounted), of C. nigritulus.

Figure 4-20. Median lobe, lateral view, of C. nigritulus.

Figure 4-21. Basal plate, ventral view, of C. nigritulus.










































Figure 4-22. U.S. states and Canadian provinces from which specimens of Cybocephalus
nigritulus have been collected.