Identification of Nipaecoccus (Hemiptera: Coccomorpha: Pseudococcidae) species in the United States, with descriptions of Nipaecoccus bromelicola sp. n. and the male of N. floridensis Beardsley Author Ellenrieder, Natalia Von Author Watson, Gillian W. Author Kinnee, Scott A. text Zootaxa 2018 2018-07-09 4444 2 163 178 journal article 29711 10.11646/zootaxa.4444.2.5 da8a0035-89c0-42db-8a5c-b154adac9900 1175-5326 1309596 4150D7BC-B075-4D2D-97E1-40CE57C2C4ED Nipaecoccus floridensis Beardsley, 2001 ( Figures 4 , 5 ) Material examined. 197 ♀ and 7 ♂. U.S.A., California: Los Angeles County, 3 ♀, Carson, in nursery on King palm, 15.vii.2013 , Salcedo leg. [CSCA, PDR # 190P06058690]; 7 ♀, Long Beach, in nursery on King palm, 9.vii.2013 , S. Ruse leg. [CSCA, PDR # 190P06058656]; 1 ♀, 1 ♂, same data but in nursery on King palm, 22.viii.2017 , C. Foy leg. [CSCA, PDR # 190P06620114]; 42 ♀, 2 ♂, Monterey Park, in nursery on Syagrus romanzoffiana and Trachycarpus fortunei , 13.xi.1995 , Papilli & Garrison leg. [CSCA, PDR # 1122913]; 10 ♀, same data but in nursery on Archontophoenix sp., 8.ix.1998 , Kendall leg. [CSCA, PDR # 1212067]; 1 ♀, same data but in nursery on Mexican fan palm, 18.viii.2005 , Walton leg. [CSCA, PDR # 1352280]; 4 ♀, North Hollywood, in nursery on King palm, 22.x.2015 , Abharian, Trung leg. [CSCA, PDR #190P06059638, DNA vouchers 15T870– 873]; 2 ♀, Santa Fe Springs, in nursery on palm, 15.xii.2005 , G. Deres leg. [CSCA, PDR # 1352477]; 6 ♀, Sun Valley, in nursery on palm, 30.x.2002 , T. Herrera leg. [CSCA, PDR # 1252840]; 11 ♀, same data but in nursery on palm, 10.ii.2015 , Abharian leg. [CSCA, PDR # 190P06059233]; 4 ♀, same data but in nursery on Phoenix roebelenii , 27.x.2015 , Abharian leg. [CSCA, PDR # 190P06059651, DNA vouchers 157950–953]; 5 ♀, Sylmar, in nursery on palm, 29.xii.2005 , R. Orsborn leg. [CSCA, PDR #c1352496]; 9 ♀, 1 ♂, same data but in nursery on Queen palm, 6.ii.2006 [CSCA, PDR # 1352580]; 7 ♀, same data but in nursery on Chamaerops humilis , 10.ii.2006 [CSCA, PDR # 1352589]; 5 ♀, same data but in nursery on Queen palm, 29.x.2015 , Abharian leg. [CSCA, PDR # 190P06059656]; 3 ♀, Westminster, in nursery on Mexican fan palm, 16.ix.1998 , Bogaert leg. [CSCA, PDR # 1212115]. Orange County, 4 ♀, Anaheim, in nursery on Syagrus romanzoffiana , 8.ix.1996 , M. Nestor leg. [CSCA, PDR # 1145089]; 8 ♀, same data but 29.vii.1997 [CSCA, PDR # 1145197]; 1 ♀, same data but 27.i.1998 [CSCA, PDR # 1204342]; 7 ♀, same data but on Washingtonia robusta , 15.x.1998 [CSCA]; 4 ♀, San Juan Capistrano, in nursery on Washingtonia sp., 15.vi.2000 [CSCA]; 4 ♀, Brea, in nursery on King palm, 2.xi.2001 , Phoutrides leg. [CSCA, PDR # 085566]. San Bernardino County, 3 ♀, 3 ♂, Chino, in nursery on Fan palm, 21.vii.2016 , Cochrane, Vigil leg. [CSCA, PDR # 360P 06202635 ]; 1 ♀, Montclair, in nursery on palm, 2.viii.2016 , Johnson leg. [CSCA, PDR # 360P06380914]; 2 ♀, Ontario, in nursery on Windmill palm, 3.i.2013 , E. Pearson leg. [CSCA, PDR # 360P06147027, DNA vouchers 13M002–003]. San Diego County, 2 ♀, Lemon Grove, in nursery on Ptychosperma elegans , 30.viii.2012 , J. Garza leg. [CSCA, PDR # 1508906, DNA vouchers 12K072–073]. Ventura County, 2 ♀, Piru, in nursery on Syagrus romanzoffiana , ant-attended, 13.iii.2012 , L. Austin leg. [CSCA, PDR # 1266501]; 4 ♀, Fillmore, in nursery, 11.iv.2000 , Evarts leg. [CSCA, PDR # 1190499]; 3 ♀, same data but on Washingtonia robusta , 15.x.2015 , J. Le leg. [CSCA, PDR # 56VP06083121, DNA voucher 15T867]; 16 ♀, same data but on Washingtonia sp., 22.x.2015 [CSCA, PDR # 56VP06083122, DNA vouchers 15T874–876]. Florida: Broward County, 1 ♀, Fort Lauderdale, intercepted at Los Angeles County, La Puente, on Phoenix roebelenii , 15.ix.2006 , M. Dennis leg. [CSCA, PDR # 1294290]; 5 ♀, Palm Bay, Helid Rd., on Syagrus romanzoffiana , 8.iv.2015 , J. Frecchete leg. [CSCA, DNA vouchers 15T527, 528, 531]. Hamilton County, 1 ♀, Jasper, in nursery on palm, 7.ii.2012 , T. Estok leg. [CSCA]. Orange County, 1 ♀, Orlando, in nursery on Butia capitata , 8.iii.2012 , A. Puppelo leg. [CSCA]. Palm Beach County, 5 ♀, Lake Worth, on Phoenix roebelenii , 4.xii.2014 , S. Simmons leg. [CSCA]. Macroscopic appearance of adult female ( Fig. 4 ). Body reddish-orange to purple in life, entirely covered by thick tufts of white to buff-colored wax. Body contents turn black in dead specimens or when placed in ethanol or KOH. Description of adult male. Slide-mounted characters ( Fig. 5 ). Body of adult male elongate oval, 1,036 ± 83 (965–1,182) µm long, width at mesothorax 310 ± 34 (272–350) µm; both hair-like (hs) and fleshy (fs) setae present on antennae and legs, only hair-like setae present on body; multilocular pores each with 4–6 loculi, but mostly with 5. Antennae about as long as half of total body length. Male macropterous, with wings each 0.8 ± 0.1 (0.6–0.9) as long as total body length and 0.4 ± 0.1 (0.3–0.5) as wide as long. Head. Width across genae 198 ± 4 (195–205) µm. Each side of head with about 9–12 dorsal head setae, 2 genal setae, no dorsal ocular setae, 1 or 2 multilocular pores and 1 or 2 discoidal pores near each scape. Ventral head setae in a group between ventral eye and ocellus and base of antenna, numbering 12–14 on each side, also 4 or 5 in a single row along longitudinal median line between ventral eyes. With 2 pairs of round simple eyes; dorsal eyes each placed approximately antero-laterally to ventral simple eyes, each 24 ± 3 (21–29) µm in diameter, ventral eyes 25 ± 2 (24–28) µm in diameter. Ocular sclerites weakly sclerotized, and weakly reticulated dorsally. Ocelli pronounced, situated laterally, each 15 ± 3 (12–19) µm in diameter, located between preocular and post-ocular ridges. Dorsal arm of midcranial ridge extending beyond posterior margin of dorsal eyes; ventral and lateral arms forming a T-shaped ridge. Preocular ridge well developed, extending postero-medially from inner margin of dorsal simple eyes. Post-ocular ridge well developed, extending from posterior to each dorsal simple eye antero-laterally past each ocellus and then fusing with the preocular ridge before extending postero-laterally to near mouth. Interocular ridge hinted at as a slightly more sclerotized bar dorsad to each ocellus. Preoral ridge well developed. Cranial apophysis slender. FIGURE 4. Nipaecoccus floridensis Beardsley. Macroscopic appearance. Adult females, immature females, crawlers and spent male cocoons on Fan palm, from a nursery in U.S.A., California, San Bernardino County. Antenna. Ten segmented and filiform; 498 ± 28 (460–544) µm long, 0.5 ± 0.1 (0.4–0.5) of total body length. Scape 35 ± 2 (30–36) µm long, with 3–5 hs. Pedicel 51 ± 4 (45–55) µm long, with 5–14 fs and 7 or 8 hs. Lengths of segments III–X (µm): III, 71 ± 6 (64–81); IV, 46 ± 8 (38–62); V, 42 ± 3 (38–62); VI, 50 ± 3 (45–52); VII, 48 ± 3 (43–50); VIII, 52 ± 3 (48–57); IX, 44 ± 4 (38–50); X, 58 ± 6 (52–69). Approximate number of bristle-shaped setae (ab), fleshy setae and hair-like setae on each of segments III–X: III 5–9 fs, 4 or 5 hs; IV 5–14 fs, 2–4 hs; V 10–13 fs, 1 or 2 hs; VI 9–12 fs, 3–5 hs; VII 9–11 fs, 4–7 hs; VIII 1 ventral ab, 6–9 fs, 4 or 5 hs; IX 1 ventral ab, 3–9 fs, 3– 7 hs; X 3 subapical ab, 3–9 fs, 3 or 4 hs, and 2 subapical capitate setae; sensilla basiconica not detected. Thorax. Prothorax. Pronotal ridge heavily sclerotized; pronotal sclerite represented by a subtriangular sclerotization dorsolaterally. Pronotum medially with, on each side, 1 or 2 setae, 4 or 5 multilocular pores, and 2 or 3 discoidal pores; post-tergite indistinct. Pronotum laterally with 0 or 1 seta, 2 multilocular pores, and 1 discoidal pore. Dorsum with 1 or 2 antero-spiracular setae, 2 antero-spiracular multilocular pores, and 1 discoidal pore; venter with 0 or 1 antero-spiracular seta, 3 antero-spiracular multilocular pores, and 0 or 1 discoidal pore. Sternum triangular, with well-developed prosternal ridge; with 0 or 1 seta, 0–2 multilocular pores, and 0 or 1 discoidal pore on each side. Anteprosternal setae absent. Mesothorax. Prescutum oval, sclerotized, prescutal ridge well developed, prescutal suture distinct, with 1–3 prescutal setae on each side. Scutum sclerotized antero-laterally, with 3–5 scutal setae on each side; prealar ridge and triangular plate well developed. Scutellum with 1 scutellar seta on each side; anterior and posterior pronotal wing processes well separated. Basisternum bounded anteriorly by a marginal ridge, posteriorly by strong precoxal ridges, median ridge absent, 7–10 basisternal setae on each side, mainly along medial area; with 1 prescutal seta on each side. Lateropleurite narrow, mesepisternum and mesepimeron distinct; furca well developed, arms divergent and extending about halfway to marginal ridge anteriorly. Mesepisternum sclerotized, subepisternal ridge long and well developed. Postalare without setae. On each side posterior to mesospiracle, with 1 or 2 postmesospiracular setae, 2–4 multilocular pores and 5–7 discoidal pores. Tegula present, with 2 or 3 tegular setae. Metathorax. Metapostnotal sclerite and metapleural ridge well developed; with 3 or 4 metatergal setae on each side; precoxal ridge well developed and metasternal apophyses distinct. Metepisternum and metepimeron weakly sclerotized; with 2 metapleural setae, 2 multilocular pores, and 2 discoidal pores posterior to each metathoracic spiracle. Metasternum membranous, on each side 1 anterior metasternal seta, 2 anterior multilocular pores, 1 anterior discoidal pore, 1 posterior metasternal seta, 1 posterior multilocular pore, and 1 posterior discoidal pore present. Wings. Hyaline, each 845 ± 50 (788–906) µm long and 326 ± 63 (256–378) µm wide. Ratio of length to width 1: 0.4 ± 0.08 (1: 0.3–1: 0.5); ratio of total body length to wing length 1: 0.8 ± 0.1 (1: 0.7–1: 0.9); alar lobe well developed and each wing with 2 or 3 alar setae and 0–2 circular sensoria. Hamulohalteres 67 ± 5 (62–76) µm long, with an apical hamulus 56 ± 8 (40–62) µm long. Legs. Metathoracic legs longest; hind femur 150 ± 8 (141–165) µm; hind tibia 191 ± 12 (169–203) µm; hind tarsus 77 ± 3 (71–79) µm; hind tarsal claw 28 ± 2 (26–31) µm. Ratio of hind femur length to hind tibia length 1: 1.3 ± 0.1 (1: 1.2–1: 1.3); ratio of hind tibia length to hind tarsus length 1: 0.4 ± 0.1 (1: 0.4–1: 0.5). Hair-like and fleshy setae present, numbering: coxa I 3–6 fs, 5–7 hs, II 3–5 fs, 5–8 hs, III 3–5 fs, 5–7 hs; trochanter I 1 or 2 fs, 2 hs, II 1– 3 fs, 2–4 hs, III 1–3 fs, 2–4 hs; femur I 4–13 fs, 5–8 hs, II 5–12 fs, 6–9 hs, III 6–17 fs, 6–9 hs; tibia I 11–17 fs, 5–8 hs, II 11–17 fs, 7–9 hs, III 12–16 fs, 7–13 hs; tarsus I 4–8 fs, 6–9 hs, II 5–8 fs, 4–7 hs, III 6–9 fs, 6–8 hs. Each trochanter with 3 campaniform sensilla arranged in a triangle medially on each side. Tibiae with apical spurs: I 2 or 3, II 3–5 , III 3–5. Tarsi each 2 segmented, tarsal digitules capitate, claw digitules acute. Abdomen. SI–VII: Small slightly sclerotized abdominal tergites present sometimes laterally on SI, SII, medially on SVI, SVII, and around base of setae on SVII. Sternites unsclerotized except for a small anterolateral area on each side of SVII. Each segment with dorsal setae numbering: SI 2 or 3; SII 2; SIII 1; SIV 1; SV 1–3; SVI 2 or 3; SVII 2–4; pleural setae and pores on each side numbering: SI 3 or 4 setae + 6–8 multilocular pores + 2 or 3 discoidal pores; SII 2 or 3 setae + 1 or 2 multilocular pores + 1 or 2 discoidal pores; SIII 2 or 3 setae + 1 or 2 multilocular pores + 1–3 discoidal pores; SIV 2 or 3 setae + 1 or 2 multilocular pores + 1–3 discoidal pores; SV 2 or 3 setae + 2 multilocular pores + 1 or 2 discoidal pores; SVI 3 or 4 setae + 2 multilocular pores + 1 or 2 discoidal pores; SVII 3–5 setae + 1–3 multilocular pores + 1 or 2 discoidal pores. Ventral setae numbering: SII 0–3; SIII 1–4; SIV 2–4; SV 3 or 4; SVI 2–4; SVII 3 or 4. Ostioles not detected. SVIII: tergite with 4 or 5 setae, 0 anteanal setae, and large medial sclerite; sternite with a small slightly sclerotized anterolateral area on each side of SVIII, without ventral setae; caudal extension rounded with 4–6 pleural setae. Glandular pouches shallow, each with 12–30 multilocular pores spreading out around pouch, 0–2 discoidal pores, 2 long glandular pouch setae, each 230 ± 28 (186–262) µm long, and 1 shorter glandular pouch seta, 89 ± 13 (71–107) µm long. Large sclerite on each of tergites IX and X. Genital segments. Penial sheath subtriangular, 103 ± 5 (95–107) µm long, 87 ± 2 (86–91) µm wide across basal ridge, ratio of penial sheath length to width 1.0: 0.9 ± 0.1 (1.0: 0.8–1.0: 0.9). Dorsum of penial sheath with 2 or 3 small setae near base of style on each side. Ventrally, style narrowing gradually to narrowly rounded apex; basal ridge well developed; lateral processes of penial sheath distinct; with 2 or 3 small setae on anterior portion of penial sheath, 0 or 1 minute setal sensillum at base of lateral process, and 0–2 on lateral process. Basal rod distinct, extending anteriorly to basal ridge. Aedeagus 68 ± 10 (52–83) µm long, approximately parallel sided, bent ventrally at a 45° angle before level of stylus and ending in a rounded tip, not extending posteriorly beyond end of penial sheath. FIGURE 5. Nipaecoccus floridensis Beardsley. Male, U.S.A., California, San Bernardino County, in nursery on Fan palm, deposited at CSCA. Enlargements of structures are not drawn to scale. TABLE 1. GenBank accession numbers, collection data and California State Collection of Arthropods (CSCA) DNA extraction codes for COI-sequenced samples of Nipaecoccus .

Taxon	GenBank accession	Collection data	CSCA DNA number
number
Nipaecoccus bromelicola	MH253682 – MH253685	U.S.A., California, Santa Barbara County, Carpinteria, in nursery on Tillandsia sp., 15.vii.2016, M. Taylor leg.,	16V450–16V453
CDFA 420P06463549
Nipaecoccus floridensis	MH290523 – MH290525	U.S.A., Florida, Broward Co., Palm Bay, Heild Rd., on Syagrus romanzoffiana , 8.iv.2015, J. Frecchete leg.,	15T527, 15T528, 15T531
Florida DPA E2015-1683-1
Nipaecoccus floridensis	MH290526 – MH290529	U.S.A., California, Los Angeles County, North Hollywood, in nursery on King palm, 22.x.2015, Abharian &	15T870–15T873
Trung leg., CDFA 190P06059638
Nipaecoccus floridensis	MH290530 – MH290533	U.S.A., California, Los Angeles County, Sun Valley, in nursery on Phoenix roebelenii , 27.x.2015, Abharian	15T950–15T953
leg., CDFA 190P06059651
Nipaecoccus floridensis	MH290534 , MH290535	U.S.A., California, Los Angeles County, Sylmar, in nursery on Queen palm, 29.x.2015, Abharian leg., CDFA	15T895, 15T896
190P06059656
Nipaecoccus floridensis	MH290536 , MH290537	U.S.A., California, San Diego County, Lemon Grove, in nursery on Seaforthia elegans , 30.viii.2012, J. Garza	12K072, 12K073
leg., CDFA 1508906
Nipaecoccus floridensis	MH290538 , MH290539	U.S.A., California, San Bernandino County, Ontario, in nursery on Windmill palm, 3.i.2013, Ed Pearson leg.,	13M002, 13M003
CDFA 360P06147027
Nipaecoccus floridensis	MH290540	U.S.A., California, Ventura County, Fillmore, in nursery on Washingtonia robusta , 15.x.2015, J. Le leg.,	15T867
CDFA 56VP06083121
Nipaecoccus floridensis	MH290541 – MH290543	U.S.A., California, Ventura County, Fillmore, in nursery on Washingtonia sp., 22.x.2015, J. Le leg., CDFA	15T874–15T876
56VP06083122
Nipaecoccus nipae complex1	MH290544	U.S.A., California, Santa Barbara County, in nursery on Washingtonia sp., Ian Swisher leg., 17.viii.2016,	16V545
(yellow wax) Nipaecoccus nipae complex1	MH290556 , MH290557	CDFA 420P06463520 U.S.A., Hawaii, Hawaii, Opihikau, on Cocos nucifera foliage, Stacy Chun leg., 3.xi.2017, HDOA 2017-394	17X976, 17X978
(buff wax) Nipaecoccus nipae complex2	MH290545 – MH290547	Ex U.S.A., Florida, Ormond Beach; intercepted at California, Contra Costa County, on Annona squamosa ,	16V483–16V485
(white wax)	Craft & Sampson leg., 27.viii.2016, CDFA 010P06556450
Nipaecoccus nipae complex3	MH290548	Ex U.S.A., Hawaii, Pahoa; intercepted at California, Los Angeles County, on cut foliage, Ogbuagu leg.,	15T449
Nipaecoccus nipae complex3	MH290549 – MH290552	9.vi.2015, CDFA 190P06059361 U.S.A., Hawaii, Kauai, Lihue Airport main terminal landscaping, on Strelitzia reginae , C. Kaneshige & L.	17X969–17X972
(yellow wax) Nipaecoccus nipae complex3	MH290553 – MH290555	Ishii leg., 6.xi.2017, HDOA 2017-392 U.S.A., Hawaii, Kauai, Lihue Airport main terminal landscaping, on Ophiopogon japonicus , E.J. Garcia & L.	17X973– 17X975
(yellow wax)	Ishii leg., 6.xi.2017, HDOA 2017-391
Nipaecoccus viridis	MH290512 – MH290521	U.S.A., Guam, Mangilao, University Campus, on Jatropha sp., L. Kikuchi leg., 16.x.2017	17X863–17X872
Nipaecoccus viridis	MH290522	Malaysia, Selangor, Klang, Jalan Kebun MARDI Station, on Artocarpus heterophylus , D. Sartiami & Hanifah	17X835
leg., 3.x.2017

Diagnosis. See comments made under N. bromelicola above. In addition, the males described as N. nipae by Afifi (1968) , from palms in South Africa (tarsal claws each 31–36 µm long), did not have the long tarsal claws which, according to Beardsley (1960 , 2001 ), characterize males of true N. nipae (42 µm); they might correspond to N. paranipae , or to another cryptic species within the N. nipae complex. Males of Afifi’s N. nipae are quite similar to the males of N. floridensis in structure and size ( Afifi 1968 ), apparently differing from the latter slightly by their smaller dorsal eyes (each 18–21 µm in diameter, versus 21.5–28.5 µm in N. floridensis ), shorter hind tarsus (each 61–67 µm, versus 71.5–78.5 µm in N. floridensis ) and hind femur (each 131–143 µm, versus 141–164 µm in N. floridensis ), and longer tarsal claw (each 31–36 µm, versus 26–31 µm in N. floridensis ). Hosts . Arecaceae : so far, recorded on Acoelorrhaphe wrightii ( Beardsley 2001 ) , Archontophoenix sp., Butia capitata , Chamaerops humilis , Phoenix roebelenii , Ptychosperma elegans , Syagrus romanzoffiana , Trachycarpus fortunei , Washingtonia robusta and Washingtonia sp. Distribution. The origin of N. floridensis , like the similar Nipaecoccus nipae , is probably Neotropical ( Beardsley 2001 ). So far, it is known only from Florida in the U.S.A. In California , it has been found only in nursery situations, where it is eradicated whenever found. Molecular data. The COI sequences of the four specimens of Nipaecoccus bromelicola analyzed are identical. For comparable sequences of COI, N. bromelicola differs from the other Nipaecoccus species sequenced so far: by 7–8% from N. floridensis ; 5.5–7.2% from members of the N. nipae complex; and by 13% from N. viridis . The COI sequences of the specimens of N. floridensis from Florida matched those of specimens of N. floridensis from California nurseries by 99–100%, and the same variability of 0–1% (out of 543–614 base pairs) was observed among the N. floridensis specimens from California . Females presenting N. nipae morphology analyzed for COI presented three distinct sequences differing by 5– 7% from each other. The COI sequences of specimens of N. viridis from Malaysia and Guam were identical. All the sequences are available at GenBank under the accession numbers and associated data indicated in Table 1 .