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VOL. 104 JANUARY 2002 NO. 1 G

EO iK (ISSN 0013-8797)

"PROCEEDINGS

of the

ENTOMOLOGICAL SOCIETY

ADAMSKI, DAVID—A new species of Glyphidocera Walsingham (Lepidoptera: Gelechioidea: Gilyphidacerdac) bison CostarRacay ! by. oko Sates ota tee A Raa eee erste ee 119

BRAILOVSKY, HARRY—A new species of Maevius Stal from Australia and some notes on the family akoyocephalidae:(Hemipteras Heteroptera) ete ncilsaoe de dane ee ae eee eben a sioner 41

BRAILOVSKY, HARRY—A new genus and a new species of Daladerini (Hemiptera: Heteroptera. Coreidac) wromiviad ag ascary wae a). pense cet Nee cee Ole eet eres ees ise BOTS: 111

BURKS, ROGER A. and JOHN D. PINTO—Reproductive and electrophoretic comparisons of Trichogramma californicum Nagaraja and Nagarkatti with the 7) minutum complex (Hyme- Hoptera: Uric hOpramiatatidae yy em aah oe te ner creat NONE eect aR tal ct tel ea aa Pr Pa 33

CARROLL, J. F—Notes on the responses of host-seeking nymphs and adults of the ticks Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae) to canine, avian, and deer-produced

SUROSTAMGE S357, co alate ais Vara a ree OL ae Raa REE RL SRY So SE CCG eal A Lee RE or) a qe) DELLAPE, P. M., M. pet C. COSCARON, and B. F AMARAL FILHO—Immature stages of Montina confusa (Stal) (Heteroptera: Reduviidae: Harpactorinae) ........................005- 168

DEWALT, R. EDWARD, DONALD W. WEBB, and AMY M. SOLI—The Neoperla clymene (Newman) complex (Plecoptera: Perlidae) in Illinois, new state records, distributions, and an LASN tit CA OMI C Ye 4e02-... lsh! cio J ae nate baie Se marae pte Rae ee neat: CRM Seas ARR an aa am 126

FERREIRA, PAULO SERGIO FIUZA and THOMAS J. HENRY—Descriptions of two new species of Fulvius Stal (Heteroptera: Miridae: Cylapinae) from Brazil, with biological and bioscorraphie notes on thes Senus {kee eres tas acs isle Mopars ese eee ears ee Meenas eee 56

GOEDEN, RICHARD D.—Descriptions of TJephritis footei and T. headricki, new species (Dip- tera: Tephritidae), with notes on their life histories in southern California .................... 142

HALL, JASON P. W.—A review of the new riodinid butterfly genus Panaropsis (Lepidoptera:

ING CorOHeRCG Ver Mtsy arava ntzted SVU) AT) Weesamtyde iat smerNey crate PeeKe A UN ents doves era ny en} Can uate ne OS 63 HARRIS, STEVEN C. and OLIVER S. FLINT, JR.—New Alisotrichia (Trichoptera: Hydroptili- dae) from Central and South America and the Greater Antilles ................0. 0... eee eee 195

(Continued on back cover)

THE

ENTOMOLOGICAL SOCIETY OF WASHINGTON

OFFICERS FOR 2002

GABRIELA CHAVARRIA, President MICHAEL G. PoGug, 7reasurer JONATHAN R. Mawps_Ley, President-Elect RonaLp A. Ocuoa, Program Chair Sruart H. McKamey, Recording Secretary STEVEN W. LINGAFELTER, Membership Chair Hotuis B. WILLIAMS, Corresponding Secretary JOHN W. Brown, Past President

Jon A. Lewis, Custodian Davip R. Smitn, Editor .

Publications Committee RAYMOND J. GAGNE THOMAS J. HENRY Wayne N. Martuis

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PROC. ENTOMOL. SOC. WASH. 104(1), 2002, pp. 1-23

REDESCRIPTION OF PARACOCCUS MARGINATUS WILLIAMS AND GRANARA DE WILLINK (HEMIPTERA: COCCOIDEA: PSEUDOCOCCIDAE), INCLUDING DESCRIPTIONS OF THE IMMATURE STAGES AND ADULT MALE

DOUGLASS R. MILLER AND GARY L. MILLER

Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Depart- ment of Agriculture, Bldg. 005, Room 137, BARC-W, Beltsville, MD 20705, U.S.A. (e-mail: dmiller@sel.barc.usda.gov; gmiller@sel.barc.usda.gov)

Abstract.—Paracoccus marginatus Williams and Granara de Willink, commonly known as the papaya mealybug was first discovered in the Caribbean in 1994 and was collected in Florida in 1998. To facilitate implementation of control measures we have prepared descriptions, illustrations, and keys for all stages of this species including: First instar, second-instar male and female, third-instar male (prepupa) and female, fourth-instar male (pupa) and female (adult), and fifth-instar male (adult). Comparisons are given between P. marginatus and other species of Paracoccus as well as other commonly encountered

mealybug species in Florida and the Caribbean.

Key Words:

mealybugs, Coccoidea, Pseudococcidae, ontogeny, pest, papaya, Carica pa-

paya, hibiscus, invasive species, pest, Florida, Caribbean

The genus Paracoccus includes some 79 species of varied distribution from _ the ‘*‘Austro-Oriental, Ethiopian, Madagasian, Nearctic, Neotropical, New Zealand, Pacif- ic, Palaearctic, and Oriental regions”’ (Ben- Dov 1994). Although most assigned species have not been recognized as major econom- ic pests, there are two notable exceptions. Paracoccus burnerae (Brain) is considered a serious pest of citrus in South Africa (Hattingh 1993) and Paracoccus margina- tus Williams and Granara de Willink has recently received attention as a pest of pa- paya and other economically important crops in the Caribbean and Florida.

Paracoccus marginatus or the papaya mealybug was originally reported from the Neotropical Region in Belize, Costa Rica, Guatemala, and Mexico (Williams and Granara de Willink 1992) and is believed to be native to at least part of Mexico and/

or Central America (Miller et al. 1999). In 1993 or 1994, it was apparently introduced into the Caribbean Islands where it spread rapidly and is considered a pest of papaya in some areas (Miller et al. 1999). It has subsequently been reported from St. Martin (Matile-Ferrero and Etienne 1996) and Guadeloupe and St-Barthelemy Islands (Matile-Ferrero and Etienne 1998). Other Caribbean distribution records include: An- tigua (Watson and Chandler 1999); Baha- mas, San Salvador Island and New Provi- dence (Watson, personal communication), Paradise Island (Meyerdirk, personal com- munication); British Virgin Islands (Watson and Chandler 1999); Cuba (Meyerdirk, per- sonal communication); Dominican Repub- lic (Meyerdirk, personal communication); Haiti (Meyerdirk, personal communica- tion); Montserrat (Watson and Chandler 1999): Nevis (Watson and Chandler 1999);

>~ROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2 Table | lo eferences for Paracoccus marginatus. #9 Reference Acacia Miller et al. (1999) Acalypha s} Miller et al. (1999)

Acalypha wilkesiana Muell.-Arg. Ambrosia cumanensis auct. non Kunth Annona squamosa L.

Bauhinia sp.

Carica papaya L.

Carica sp.

Cestrum nocturnum L.

Citrux x paradisi Macfad. (pro sp.) Clerodendrum paniculatum L. Coccoloba sp.

Fistulosa sp.

Guazuma ulmifolia Lam.

Hamelia patens Jacq.

Hamelia sp.

Hibiscus sp.

Hibiscus rosa-sinensis L.

Ipomoea carnea Jacq.

Ipomoea sp.

Jatropha integerrima Jacq. Jatropha sp.

Malvaviscus penduliflorus DC.

Hamon (personal communication) Williams and Granara de Willink (1992) Miller et al. (1999)

Hamon (personal communication) Hamon (personal communication) Williams and Granara de Willink (1992) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Miller et al. (1999)

Hamon (personal communication) Miller et al. (1999)

Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Hamon (personal communication) Miller et al. (1999)

Hamon (personal communication) Hamon (personal communication) Hamon (personal communication)

Manihot chlorosticta Standl. & Goldman Manihot esculenta Crantz Mimosa pigra L.

Williams and Granara de Willink (1992), Miller et al. (1999) Williams and Granara de Willink (1992), Miller et al. (1999) Williams and Granara de Willink (1992)

Williams and Granara de Willink (1992), Miller et al. (1999)

Parthenium hysterophorus L.

Persea americana P. Mill.

Plumeria rubra L.

Plumeria sp.

Rhaphiolepis umbellata (Thunb.) Makino Sida sp.

Solanum melongena L.

Uniola paniculata L.

Zea mays L.

Miller et al. (1999)

Hamon (personal communication) Hamon (personal communication) Hamon (personal communication)

Williams and Granara de Willink (1992), Miller et al. (1999)

Miller et al. (1999) Hamon (personal communication) Miller et al. (1999)

Puerto Rico (Meyerdirk, personal commu- nication); St. Barthelemy (Meyerdirk, per- sonal communication); St. Kitts (Watson and Chandler 1999); and U.S. Virgin Is- lands (Watson and Chandler 1999). Addi- tionally, P. marginatus has recently been reported from French Guiana, South Amer- ica (Matile-Ferrero et al. 2000). Paracoccus marginatus was first discovered in the Unit- ed States in Manatee and Palm Beach coun- ties, Florida in 1998 and has since been col- lected in Alachua, Brevard, Broward, Col- lier, Dade, Hillsborough, Manatee, Martin, Monroe, Palm Beach, Pinellas, Polk, Sara- sota, and Volusia counties (Hamon, person-

al communication). Since its introduction into Florida, it has been collected 80 times in 30 different cities on 18 species of hosts. The species has apparently spread rapidly in the state.

Paracoccus marginatus has been report- ed from more than 25 genera of host plants including economically important crops such as papaya, citrus, yams, cassava, and hibiscus. Papaya mealybug hosts along with associated references are included in Table 1. Although this mealybug has also been recorded from many other hosts, these have yet to be confirmed and are not included in Wable ws

VOLUME 104, NUMBER 1

Williams and Granara de Willink (1992) published the first taxonomic paper on Par- acoccus marginatus and included a descrip- tion and illustration of the adult female. They also provided descriptions and a key for the 21 species of Paracoccus occurring in Central and South America. The purpose of this research is to describe and illustrate all instars (including the adult male and fe- male) of this potentially serious invasive species in order to facilitate its recognition as part of any control initiatives that may be undertaken.

METHODS

Terminology in the descriptions follows that of Williams and Granara de Willink (1992) and Gimpel and Miller (1996) for adult females and immatures and that of Af- ifi (1968) for adult males. Measurements were made with an ocular micrometer using a Leica DMRB compound microscope. Numbers and measurements are from 10 specimens and are given as an average fol- lowed by the range in parentheses. Asso- ciated enlargements of various structures on the illustrations are not proportional. All specimens studied are from the National Museum of Natural History, Coccoidea Collection, Beltsville, MD. Information list- ed in the SPECIMENS EXAMINED section is verbatim from information recorded on the microscope slides. The abbreviation ‘‘ad.”’ refers to adult specimens.

Tables 3—6 contain 12 of the most com- mon mealybugs in the Caribbean region and were selected for comparison to P. marginatus. Included species are: Dysmi- coccus brevipes (Cockerell); Ferrisia vir- gata (Cockerell); Maconellicoccus hirsutus (Green); Nipaecoccus nipae (Maskell); Phenacoccus gossypii Townsend and Cock- erell; Phenacoccus madeirensis Green; Phenacoccus solenopsis Tinsley; Planococ- cus citri (Risso); Planococcus minor (Mas- kell); Pseudococcus jackbeardsleyi Gimpel and Miller; Pseudococcus longispinus (TYar- gioni Tozzetti); and Pseudococcus viburni (Signoret). Phenacoccus gossypii and P.

madeirensis are listed together as are Plan- ococcus citri and P. minor because of mor- phological similarities. There are other spe- cies of Phenacoccus, Pseudococcus, and Nipaecoccus that could be included in this list, but they are also so similar in general appearance to other species in the genus that characters of one species in the genus are sufficient to distinguish all species in the immature instars.

Host plant names included in Table 1 were verified in Integrated Taxonomic In- formation System (Anonymous 2001).

RESULTS

Paracoccus marginatus Williams and Granara de Willink

Suggested Common Name: Papaya mealybug

(Figs. 1-8) KEY TO INSTARS ile Without wings or wing buds on thorax ... 4 - With wings or wing buds on thorax ..... 2

2(1). Genitalia weakly sclerotized, aedeagus ab- sent; wings represented by buds less than 2 length of body, without a vein ......... 3

~ Genitalia heavily sclerotized, aedeagus ap-

parent; wings approximately as long as body, with small basal vein (Fig. 8) ..... Os PRES el! Pie Od at S| Ac. Oe ERR ME Se OREM C adult male

. Antenna unsegmented; wing buds small,

less than length of thorax; head without sclerotization (Fig. 6) prepupal male

- Antenna with 10 segments; wing buds large,

about same length as thorax; head with weak sclerotization (Fig. 7) ..... pupal male

. Antennae 6-segmented

Antennae with 7 or 8 segments

. Without tubular ducts or with less than 5

SUCHIGUGISE Sin ces o = ces = Sr etecre.s 6

With more than 5 tubular ducts particularly

onidorsalfabdomens(Bi1e7s5)) tan tee second-instar male

Third antennal segment with more than 4

setae; anal-lobe cerarius with at least 1 aux-

iliary seta

- Third antennal segment with 4 setae; anal-

lobe cerarius without auxiliary setae (Fig. A) ccoreg cy. s Venchep acy Pye Rags Gycatebye Paes first instar

. Third antennal segment with 5 setae; tibia

divided by tarsus 0.9(0.8—1.0) (Fig. 3) easter suse ee Rees eee ee second-instar female - Third antennal segment with 9 setae; tibia

ey) NM ay

6(5).

4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

divided by ta 1.2-1.4) (Fig. 2) ... Te third-instar female (in part) 8(4). Antenna nted; with less than 5 mul- tilocular pores; without vulva (Fig. 2) . third-instar female (in part) Antenna 8-segmented; with more than 5

multilocular pores; with vulva (Fig. 1) ... adult female

Type data.—We have not examined type material of this species, but have studied many of the specimens that were used in the original description but were not in- cluded in the type series.

Etymology.—The species epithet is de- rived from a Latin word meaning enclosed within a border and refers to the border of oral rim tubular ducts (Williams and Gran- ara de Willink 1992).

ADULT FEMALE (Fig. 1)

Field features.—Body yellow, dusted with mealy wax not thick enough to hide body color, without discrete bare areas on dorsum, with many short waxy filaments around body margin. Ovisac developed be- neath and behind adult female.

Slide-mounted characters—Body 2.2(1.5— 2.7) mm long, 1.4(0.9—1.7) mm wide.

Dorsum with 16(14—17) pairs of cerarii; ceramml. 2.4.5. 7.-and) 9 wath 2 conical setae (Fig. 1M); cerarii 3, 6, and 16 with 3(2—3) conical setae; cerarii 8, 11, and 17 with 2(0—2); cerarii 10 and 14 with 1(0—2) conical setae; cerarii 12, 13, and 15 with 2(0—3) conical setae. Cerarius 12 without auxiliary setae, with 2(0—3) conical setae, 5(0—8) trilocular pores, 1(0—3) discoidal pores. Anal-lobe cerarius (Fig. 11) with I(1—3) auxiliary setae (Fig. 1J), 2 conical setae, 13(10—18) trilocular pores Gigs i€); 2(0—3) discoidal pores (Fig. 1K). Dorsal body setae (Fig. 1L) more slender than cer- arian setae. Multilocular pores absent; tri- locular pores scattered over surface, most abundant near setae; discoidal pores rare, about 2 diameter of trilocular pore. Oral- rim tubular ducts (Fig. 1B) usually restrict- ed to marginal areas associated with cerarii, | specimen examined with | mediolateral

duct on segment I and | in medial area of mesothorax; of 21 specimens examined cer- arius | without associated oral rim, cerarius 2 with associated oral rim in 20 of 21 spec- imens, cerarius 3 with 4 on 21 specimens, cerarius 4 with 12 on 21, cerarius 5 with 11 on 21, cerarius 6 with 17 on 21, cerarius 7 with 18 on 21, cerarius 8 with 20 on 21, cerarii 9 and 10 without associated oral rims, cerarius 11 with 15 on 21, cerarius 12 with 4 on 21, cerarii 13, 15, and 16 without associated oral rim, cerarius 14 with 2 on 21, and cerarius 17 with 15 oral rims on 21 specimens. Oral-collar tubular ducts absent. Longest submedial seta on segment VII 10(8—14) p long; 1(0—2) submedial setae on segment VIII, when present longest seta 13(8—18) p long.

Anal-ring seta. 136(120—150) wp long; 1.4(1.2—1.7) times as long as width of anal ring.

Venter with multilocular pores (Fig. 1G) usually in posterior and anterior bands on segments VI—-VIII and restricted to poste- rior band on segments IV and V, | or 2 specimens with | or 2 pores on segment III or with a few pores on anterior margins of segments IV and V; 3 of 10 specimens with 1 multilocular pore near base of front or hind leg. Trilocular pores concentrated near setal bases. Discoidal pores uncommon, of same size as on dorsum. Oral-rim tubular ducts in mediolateral areas from prothorax to segment I, with 4(3—6) ducts on each side of body. Oral-collar tubular ducts of 1 size, IN Conspicuous marginal clusters along body margin from segments II-VI, often with 2 or 3 pores on segment I, also present in medial and mediolateral areas of abdom- inal segments HII—VIII, present on thorax in seta clusters near hind 2 pairs of legs, oc- casionally with | or 2 along body margin of thorax especially in area laterad of an- terior spiracle and front legs, absent from head. Setae as follows: 4 cisanal (Fig. 1H), longest 52(45—68) pw long; longest anal-lobe seta 170(155—200) w long; longest seta on trochanter 104(95—110) p long. Anal-lobe

VOLUME 104, NUMBER |

Fig. 1. Adult female of Paracoccus marginatus. A, Detail of front leg. B, Oral-rim tubular duct. C, Trilocular pore. D, Translucent pores. E, Oral-collar tubular duct. E Detail of hind leg. G, Multilocular pore. H, Cisanal seta. I, Anal-lobe cerarius. J, Auxiliary seta. K, Discoidal pore. L, Dorsal body seta. M, Cerarian seta.

6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bar conspicuoi ler than base of anal bar set

Circulu 80) wp wide, generally di- vided b segmental line. Labium 137(125—i162) » long. Antenna 8-segment-

ed, 372(335—400) pw long. Legs with trans- lucent pores restricted to hind coxa, ventral surface (when leg is lying flat as shown in illustration) with 40(14—62) pores, dorsal surface (Fig. 1D) with 79(54—108) pores. Hind femur 209(195—225) w long; hind tib- ia 211(185—228) pw long; hind tarsus 94(91— 100) » long. Hind tibia/tarsus 2.2(2.0—2.3); hind femur/tarsus 1.0(0.9—1.1). Length of hind femur divided by greatest width of hind femur 3.7(3.0—4.2). Hind tibia with 15(14—19) setae. Claw digitules on all legs clubbed, approximately same size. Tarsal digitules (Fig. 1F) on hind 2 pairs of legs clubbed, each tarsus with 1| digitule notice- ably longer and with club slightly larger than other; tarsal digitules (Fig. 1A) on front pair of legs of 2 different sizes and shapes, | digitule on each tarsus clubbed and robust, other digitule without club, slender.

Notes.—The above description is based on 240 specimens from 41 localities. The adult female can be distinguished from all other instars by having multilocular pores, translucent pores on hind coxa, and a vulva.

THIRD-INSTAR FEMALE (Fig. 2)

Slide-mounted characters—Body 1.1(0.7— 1.8) mm long, 0.7(0.3—1.1) mm wide.

Dorsum with 6(1—10) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius with 1(1—2) auxiliary setae, 2 con- ical setae, 5(4—7) trilocular pores, 0(O0—1) discoidal pores. Dorsal body setae more slender than setae. Multilocular pores absent; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular duct rarely present near position of cerarius 8 (of 10 specimens

cerarian

examined, 4 had | oral rim or large oral collar on at least one side of body). Oral- collar tubular ducts absent. Longest sub- medial seta on segment VII 7(5—10) p long; 1(0—2) submedial setae on segment VIII, when present longest seta 7(5—9) p long.

Anal-ring seta 87(78-92) wp long; 1.6(1.3—1.8) times as long as width of anal ring.

Venter with multilocular pores absent (present on | of 10 specimens examined, restricted to segment VIII). Trilocular pores concentrated near setal bases. Discoidal pores uncommon, of same size as on dor- sum. Oral-rim tubular ducts sometimes pre- sent near body margin on abdominal seg- ment II, III, or on metathorax (of 10 spec- imens examined, 4 had 1 or 2 oral rims or large oral collar on at least one side of body). Oral-collar tubular ducts absent. Se- tae as follows: 4 cisanal, longest 33(22—50) w long; longest anal-lobe seta 132(112— 158) pw long; longest seta on trochanter 63(50—-85) ww long. Anal-lobe bar slightly narrower than on adult female.

Circulus 47(22—80) w wide, generally di- vided by intersegmental line. Labium 85(70—-100) w long. Antenna 6- or 7-seg- mented (of 10 specimens examined 5 had 6-segments with weak indication of partial division between segments 3 and 4), when 6-segmented, segment 3 with 9 setae, 233(205—262) pw long. Legs without trans- lucent pores. Hind femur 114(100—138) p long; tibia 108(98—-127) w long; tarsus 84(80-90) pw long. Hind tibia/tarsus L2C1.2=14): hind femiur/tarsus 121 G@e0= 1.2). Length of hind femur divided by greatest width of hind femur 2.5(2.0—3.1). Hind tibia with 9(7—10) setae. Claw and tar- sal digitules same as on adult female except sometimes | claw digitule slightly smaller that other.

Notes.—The above description is based on 42 specimens from 12 localities. The third-instar female can be distinguished from all other instars by having 6- or 7-seg- mented antennae, when 6-segmented with

VOLUME 104, NUMBER 1

Fig. 2. Third-instar female of Paracoccus marginatus.

8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

hind tibia divided by hind tarsus length usu- ally 1.2. and about 9 setae on the hind tibia. SECOND-INSTAR FEMALE (Fig. 3)

Field features.—Body color yellow.

Slide-mounted characters ——Body 0.7(0.5— 0.8) mm long, 0.4(0.3—0.5) mm wide.

Dorsum with 6(4—11) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius with | auxiliary setae, 2 conical se- tae, 2(2—3) trilocular pores, sometimes with 1 discoidal pore. Dorsal body setae more slender than cerarian setae. Multilocular pores absent; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular ducts absent. Oral- collar tubular ducts absent. Longest sub- medial seta on segment VII 6(5—6) pw long; without submedial setae on segment VIII.

Anal-ring seta 60(52-70) w long; 1.5(1.3-1.8) times as long as width of anal ring.

Venter with multilocular pores absent. Trilocular pores concentrated near setal ba- ses, absent in medial area of abdomen. Dis- coidal pores rare. Oral-rim tubular ducts ab- sent. Oral-collar tubular ducts absent. Setae as follows: 4 cisanal, longest 20(18—25) long; longest anal-lobe seta 95(88—102) w long; longest seta on trochanter 49(42—55) w long. Anal-lobe bar narrower than on adult female.

Circulus 40(20—75) j wide, generally di- vided by intersegmental line. Labium 69(62—75) w long. Antenna 6-segmented, 173(152—185) pw long; antennal segment 3 with 5 setae. Legs without translucent pores. Hind femur 78(72—82) w long; hind tibia 66(58—72) ww long; hind tarsus 69(65— 72) w long. Hind tibia/tarsus 0.9(0.8—1.0); hind femur/tarsus 1.2(1.1—-1.3). Length of hind femur divided by greatest width of hind femur 2.3(2.1—2.5). Hind tibia with 9(8—9) setae. Claw and tarsal digitules same

as on adult female except | claw digitule conspicuously smaller on all legs.

Notes.—The above description is based on 64 specimens from 9 localities. The sec- ond-instar female can be distinguished from all other instars by lacking oral-collar tu- bular ducts and multilocular pores, and by having 5 setae on the third antennal seg- ment.

First INSTAR (GENDER NOT DETERMINED) (Fig. 4)

Slide-mounted characters.—Body 0.4(0.3— 0.5) mm long, 0.2(0.2—0.3) mm wide.

Dorsum with 9(7—10) pairs of cerarii; cerarii indefinite, when present, with 2 con- ical setae and | trilocular pore between con- ical setae. Cerarius 12 absent. Anal-lobe cerarius without auxiliary setae, 2 conical setae, | trilocular pore, without discoidal pores. Dorsal body setae more slender than cerarian setae. Multilocular pores absent; trilocular pores scattered over surface, forming 2 longitudinal lines on each side of abdomen, excluding cerariian setae. Discoi- dal pores absent. Oral-rim tubular ducts ab- sent. Oral-collar tubular ducts absent. Lon- gest submedial seta on segment VII 5(4—8) w long; without submedial setae on segment VUl.

Anal-ring seta 37(30—42) w long; 1.4(1.2—1.6) times as long as width of anal ring.

Venter without multilocular pores. Tri- locular pores restricted to 10 positions on each side of body on head, thorax, and an- terior abdomen (see Fig. 4 for position la- bels); triloculars in positions | to 7 and 10 rarely absent, pores in positions 8 and 9 of- ten absent. Discoidal pores in submarginal line on each side of abdomen, usually with | pore posterior of each spiracle. Oral-rim and oral-collar tubular ducts absent. Setae as follows: 4 cisanal, longest 17(12—22) w long; longest anal-lobe seta 58(45—69) w long; longest seta on trochanter 38(35—42) w long. Anal-lobe bar narrower than on adult female.

Circulus 37(20—55) wide, generally di-

VOLUME 104, NUMBER 1

Fig. 3. Second-instar female of Paracoccus marginatus.

9

10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

, 1ret—i Pi, aa) > Paes . : : = San = Fig. 4. First-instar of Paracoccus marginatus, gender not determined. Numbers refer to positions of ventral trilocular pores.

VOLUME 104, NUMBER 1

vided by intersegmental line. Labium 48(44—-52) ww long. Antenna 6-segmented, 132(120—148) ww long; antennal segment 3 with 4 setae. Legs without translucent pores. Hind femur 59(55—62) pw long; hind tibia 50(45—55) pw long; hind tarsus 57(53— 62) w long. Hind tibia/tarsus 0.9(0.8—0.9); hind femur/tarsus 1.2(1.1—1.3). Length of hind femur divided by greatest width of hind femur 2.3(1.8—2.5). Hind tibia with 9 setae. Claw and tarsal digitules same as on adult female except | claw digitule con- spicuously smaller on all legs.

Notes.—The above description is based on 86 specimens from 6 localities. The first- instar can be distinguished from all other instars by having no auxiliary setae in the anal-lobe cerarius, 2 longitudinal lines of trilocular pores on dorsal abdomen on each side of body (excluding cerariian setae), and antennal segment 3 with 4 setae.

SECOND-INSTAR MALE (Fig. 5)

Field features.—Body color usually pink, occasionally yellow.

Slide-mounted characters.—Body 0.6(0.5— 1.0) mm long, 0.3(0.2—0.6) mm wide.

Dorsum with 4(2—5) pairs of cerarii; cer- arii indefinite, when present, with 2 conical setae and | trilocular pore between conical setae. Cerarius 12 absent. Anal-lobe cerar- ius with 1(1—2) auxiliary setae, 2 conical setae, 2(2—3) trilocular pores, without dis- coidal pores. Dorsal body setae more slen- der than cerarian setae. With 1(0—2) multi- locular pores in medial areas of thorax and/ or head, present on 6 of 10 specimens ex- amined; trilocular pores scattered over surface, most abundant near setae; discoidal pores rare, about % diameter of trilocular pore. Oral-rim tubular ducts absent. Oral- collar tubular ducts abundant over surface, of 1 size. Longest submedial seta on seg- ment VII 6(5—8) long; without submedial setae on segment VIII.

Anal-ring seta 54(48—-58) w_ long; 1.3(1.2-1.5) times as long as width of anal ring.

11

Venter with multilocular pores mesad of each pair of legs, for hind pair of legs lo- cated on segment III, with 4(2—5) pores on body. Trilocular pores concentrated near se- tal bases. Discoidal pores rare. Oral-rim tu- bular ducts absent. Oral-collar tubular ducts of 2 sizes: larger size same as on dorsum, located marginally; smaller size present in longitudinal line along submargin of abdo- men. Setae as follows: 4 cisanal, longest 21(17—26) pw long; longest anal-lobe seta 91(83-108) ww long; longest seta on tro- chanter 47(40—55) pw long. Anal-lobe bar narrower than on adult female.

Circulus 50(40—75) w wide, generally di- vided by intersegmental line. Labium 67(62—72) w long. Antenna 6-segmented, 171(160—188) ww long; antennal segment 3 with 5 setae. Legs without translucent pores. Hind femur 82(75—92) long; hind tibia 68(61—80) pw long; hind tarsus 62(52— 70) w long. Hind tibia/tarsus 1.1(1.0—1.2); hind femur/tarsus 1.2(1.1—1.4). Length of hind femur divided by greatest width of hind femur 2.6(2.2—3.0). Hind tibia with 9(8—9) setae. Claw and tarsal digitules same as on adult female except | claw digitule conspicuously smaller on all legs.

Notes.—The above description is based on 86 specimens from 7 localities. The sec- ond-instar male can be distinguished from all other instars by having dorsal oral-collar tubular ducts and multilocular pores near | or more pairs of legs. In addition, live and alcohol preserved specimens tend to appear more narrowly elongate than other associ- ated second-instar females, third-instar fe- males, or first instars.

Both field collected and laboratory reared early instars of P. marginatus exhibit pink and yellow forms. We hypothesize that the pink forms are males. Pink forms were col- lected in the field on St. Kitts, W.I. In ad- dition, Richard Warkentin (USDA, APHIS- PPQ, Biological Control Technical Special- ist) collected and sorted a vial of pink spec- imens and a vial of yellow ones from the Papaya mealybug rearing facilities in St. Thomas, U.S.V.I. However, the results from

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leis 2)

Second-instar male of Paracoccus marginatus.

VOLUME 104, NUMBER 1

both collection sites were inconclusive. Al- though field collected pink forms were mostly second instar males, there were a few second instar females present. Results of the laboratory reared specimens were also inconclusive. While specimens of the pink form yielded only second instar males, specimens of the yellow form yielded both second instar males and second instar fe- males although there were many more fe- males than males. Contamination, especial- ly for field collected specimens may be a possibility for the discrepancies but, this is probably unlikely for the specimens col- lected at the rearing facilities.

THIRD-INSTAR MALE (PREPUPA) (Fig. 6)

Slide-mounted characters.—Body 0.9(0.8— 1.1) mm long, 0.4(0.3—0.4) mm wide.

Dorsum without cerarii; posterolateral margins of segments V, or VI, VII, and VIII each with 2 setae conspicuously longer than remaining setae on abdominal segments. Multilocular pores in medial areas of head, forming row on prothorax and metathorax, usually without pores on mesothorax, oc- casionally with | or 2 medially, in rows on most abdominal segments, fewer in medial area, absent from segments VIII and IX; tri- locular pores absent; discoidal pores rare. Oral-rim tubular ducts absent. Oral-collar tubular ducts present around body margin, medial and submedial ducts sometimes pre- sent on prothorax, metathorax and 1 or 2 abdominal segments. Longest submedial seta on segment VII 18(15—20) wp long; without submedial setae on segment VIII.

Anal-ring setae absent; anal ring 25(20— 28) ww wide.

Venter with multilocular pores near an- terior margin on head, near spiracles, legs, and in medial areas of pro- and mesothorax on thorax, in rows on abdominal segments, sparse medially, absent from segments VIII and IX. Trilocular pores absent. Discoidal pore located near each pair of legs. Oral- rim tubular ducts absent. Oral-collar tubular

13

ducts restricted to margin. Longest anal- lobe seta 67(50—78) wp long.

Circulus appearing collapsed, 62(45—98) ut wide, resting on intersegmental line. La- bium absent. Antennal segments indistinct, 226(215—248) pw long. Hind femur 90(82— 95) » long; division between hind tibia and tarsus indistinct, hind tibia + tarsus 134(130—140) p long. Length of hind fe- mur divided by greatest width of hind fe- mur 2.7(2.2—3.0). Wing buds of mesothorax protruding from lateral margin, 67(50—78) w long. Hamulohalterae represented by small swelling on lateral margin of meta- thorax.

Notes.—The above description is based on 16 specimens from 3 localities. The pre- pupa can be distinguished from all other in- stars by having multilocular pores, oral-col- lar tubular ducts, antennae without definite segmentation, tibia + tarsus fused, no la- bium, no aedeagus, and no definite constric- tion for the head.

FOURTH-INSTAR MALE (PUPA) (Fig. 7)

Slide-mounted characters.—Body 1.0(0.9— 1.0) mm long, 0.3(0.3—0.4) mm wide.

Dorsum without cerarii; posterolateral margins of segments III, IV, or V, to seg- ment VIII each with 2 setae conspicuously longer than remaining setae on abdominal segments. Multilocular pores absent from head, forming conspicuous row on protho- rax, mediolateral cluster on metathorax, without pores on mesothorax, in mediolat- eral clusters on each side of abdominal seg- ments I-VI or VII; trilocular pores absent; discoidal pores associated with multilocu- lars and oral collars. Oral-rim tubular ducts absent. Oral-collar tubular ducts present near body margin of prothorax and abdom- inal segments I or II to VII or VII, forming clusters of 2(1—5) ducts. Longest submedial seta on segment VII 20(16—28) wp long; without submedial setae on segment VIII.

Anal-ring setae absent; anal ring 27(25— 30) » wide.

Venter with multilocular pores absent

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 6.

Third-instar male or prepupa of Paracoccus marginatus.

VOLUME 104, NUMBER 1

Fig. 7. Fourth-instar male or pupa of Paracoccus marginatus.

16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

from head. row of pores between front cox-

mainder of thorax near spi-

ae, present on rr

racles and legs, in mediolateral clusters of 2(1—4) pores on each side of segments H— VI. VII, or VIL. Trilocular pores absent.

Discoidal pores associated with oral-collars and multiloculars. Oral-rim tubular ducts absent. Oral-collar tubular ducts present near body margin of prothorax usually forming cluster of several ducts, sometimes absent from abdomen or with | duct near body margin on each of abdominal seg- ments II-VII or VIII. Longest anal-lobe seta 60(48—72) pw long.

Circulus ill-defined. Labium absent. An- tenna 10-segmented, 357(345—375) w long. Hind femur 109(105—115) p long; hind tib- ia 111(105—112) p long; hind tarsus 80(75— 88) » long. Hind tibia/tarsus 1.4(1.3—1.5); hind femur/tarsus 1.0. Length of hind femur divided by greatest width of hind femur 3.3(3.2—3.4). Wing buds of mesothorax pro- truding from lateral margin, 336(250—385) w long. Hamulohalterae 36(22—42) w long.

Notes.—The above description is based on 8 specimens from | locality. The pupa can be distinguished from all other instars by having multilocular pores, oral-collar tu- bular ducts, 10 segmented antennae, no la- bium, no aedeagus, and a slight constriction between the thorax and head.

ADULT MALE (Fig. 8)

Slide-mounted characters.—Body elon- gate oval, 1.0(0.9—1.1) mm long; greatest width at thorax 0.3(0.2—0.3) mm.

Dorsum with | pair of tail-forming pore clusters; each cluster with 2 elongate setae approximately 250 wp long, | sometimes 2 additional shorter setae, 38(34—42) multi- locular pores, and | or 2 discoidal pores. Multilocular pores in marginal areas of pro- thorax and each abdominal segment, with 5(3-7) on each side of segment I, 2(1—3) on each side of segment II, 2(1—2) on III, 2(1—3) on IV, 2(1—3) on V, 2(1-3) on VI, 1(1—2) on each side of segment VII, mul- tilocular pores with 4 or 5 loculi, quinquel-

oculars predominate (Fig. 8G); normally without pores on head (1 of 10 specimens with | pore near lateral arm of midcranial ridge). Discoidal pores (Fig. 8F) associated with lateral abdominal multilocular pores. Body setae bristle shaped. Small abdominal tergites present on mid-dorsum of segments I-III and dorsum of segment VIII. Dorsal abdominal tergites usually without associ- ated setae. Metapostnotal ridge conspicu- ous. Scutellum rectangular, with several medial setae. Scutum sclerotized through- out except for a median longitudinal clear area which bears several setae. Prescutum rectangular with well defined prescutal ridge, weakly defined prescutal suture and several setae. Pronotal ridge heavily scler- otized. Hamulohalterae 75(67—82) w long, with 1 apical hooked seta. Mesothoracic wings 932(889—988) p long, each with 2—3 basal setae. Head width 180(148—193) p; dorsal eye 34(30—40) p in diameter, lateral ocellus 17(12—20) wp in diameter and locat- ed at junction of preocular and postocular ridges. Dorsal arm of midcranial ridge (Fig. 8A) extending beyond posterior margin of dorsal eye. Median crest weakly sclerotized with several setae. Ocular sclerite weakly sclerotized.

Penial sheath (Fig. 8C and 8E) 95(86— 106) pw long, 70(62—74) pw wide with distinct ventral lobes; length/width ratio 1.4. Ae- deagus (Fig. 8D) 68(54—79) w long, broad and apically truncate.

Venter with hair-like setae only, present medially, submedially and laterally of most abdominal segments as well as few scat- tered prosternal and basisternal setae. Ab- dominal sclerotization confined to segment VIII. Prosternal ridge well developed, ster- nite weakly sclerotized. Preoral ridge weak- ly developed. Ocular sclerite weakly scler- otized near ventral eye. Ventral midcranial ridge well developed, with lateral arms. Ventral eye 39(32—44) w in diameter.

Antenna 10-segmented with bristle- shaped and fleshy setae, capitate setae pre- sent on apical segment; segments I 37(30— 40) w long (Fig. 8H); I 55(49—-62) pw long

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Fig. 8. Adult male of Paracoccus marginatus. A, Frontal view of midcranial ridge. B, Detail of front leg. C, Ventral view of penial sheath. D, Aedeagus. E, Lateral view of penial sheath. F Discoidal pore. G, Quin- quelocular pore. H, Detail of scape and pedicel. I, Detail of apical segment.

18 ROCEREDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2 omparisons of New World Paracoccus. Bold indicates characters that differ from Paraco fare fee Marginal Cluster Oral Collars Number of Near Anterior Pores on

Par Circulus Cerarii Oral Rim Distribution Spiracle Hind Tibiae marginatus yes 16-17 margin only no no alazanensis yes 17 margin and abdomen no yes ascius yes IL7/ scattered yes yes baccharidicola yes a scattered yes yes circuliprivis no 7 margin and thorax no yes decorus no 7-9 throughout except head no yes ferrisi variable 12-17 usually margin and abdomen yes yes hamoni yes 12 scattered no yes herreni yes 16 scattered yes yes juniperi yes 5-17 scattered yes yes lycopersici yes 4-7 scattered yes yes mexicanus yes 15-17 scattered no yes myrtacearum no 14 scattered no yes oneratus yes 5 many, scattered no yes ordinis yes 15-16 scattered yes yes reductus yes 0 absent no yes salviacola yes 16 scattered yes yes solani variable 6-7 scattered no yes townsendi yes 13 absent yes no turrialbensis no wT abdomen no yes villanuevai yes 7 scattered yes no

(Fig. 8H); III 68(62—74) wp long; VI 52(45— 57) w long; VII 53(47-59) pw long; VIII 53(47—-59) w long; IX 46(37—49) pw long; X 46(49-69) w long (Fig. 81); total length 534(469-563) pw long. Hind femur 145(133—148) p long; hind tibia 199(163- 222) » long; hind tarsus 83 (69-86) yp long; hind tarsal claw 27(22—32) w long; hind fe- mur/tibia ratio 0.7; hind tibia/tarsus ratio 2.4(2.3—2.5); leg setae bristle shaped. Tarsal digitules capitate; claw digitules acute (Fig. 8B).

Notes.—The above description is based on 64 specimens from 8 localities. The adult male can be distinguished from all other instars by having a distinct aedeagus, lateral pore clusters, a heavily sclerotized thorax and head, and by having well-de- veloped wings.

SPECIMENS EXAMINED

UNITED STATES: Florida: Broward County, Ft. Lauderdale, on Hibiscus sp.,

5-XI-1998, by W. Thiel. Manatee County, Bradenton, on H. rosa-sinensis, 24-VII- 1998, by W. Clifton (2 ad. 2). Palm Beach County, Boca Raton, on Hibiscus rosa-si- nensis, 1998; by J. Lofquist (l\;adeeg): BRITISH VIRGIN ISLANDS: Great Ca- manoe Island, on /pomoea sp., 16-X-1996, by R. E Denno (7 ad. 2°). DOMINICAN REPUBLIC: Locality unknown, on Persea americana, 19-VII-1994, by J. Sanchez (1 ad. 2). MEXICO: Baja California—La Paz, on Carica papaya, 13-1X-1978, by G. Bux- ton (16 ad. 2, 3 third-instar 2, 3 second- instar d). Colima—Cofradia de Juarez, on Carica papaya, 8-V1-1999, by H. Gonza- lez, J.