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Acanthococcus torikurai Tanaka sp. nov. Proposed Japanese common name: Tsutsuji-fukuro-kaigaramushi. Material examined. Holotype: JAPAN: / Hokkaido, Tokoro-gun, / Kunneppu-cho, / Rhododendron sp., / 5. vi.1990, / coll. H. Torikura: adult female mounted singly (ELKU). Paratypes: same data as for holotype: 7 adult females mounted singly (3 ELKU, 4 EUMJ). Description (n = 8) Live adult female: not seen. Slide-mounted adult female (Fig. 1): body oval, 2.5 (2.0–2.6) mm long, 1.8 (1.2–1.8) mm wide. Antenna 7 segmented, 282–289 (253–298) µm long; segment lengths in µm: I: 55–56 (40–60), II: 37–48 (30–48), III: 50–52 (43–57), IV: 46–56 (40–58), V: 21–24 (19–26), VI: 18–20 (15–22), VII: 41 (38–43); segments III and V without flagellate setae; other segments each with a few flagellate setae; apical segment with apical setae 43–52 (43–65) µm long, also with 2 or 3 (mostly 3) sensory falcate setae, longest falcate setae 32–40 (32–45) µm long; 2 preapical segments also each with 1 sensory falcate seta; falcate setae length on segment VI, 31–30 (20–45); V, 28–29 (28–43). Frontal lobe present, 29–30 µm (25–60 µm) wide. Frontal tubercle absent. Eyes each situated on venter near margin. Anterior spiracles each 27–28 (21–29) µm in diameter; posterior spiracles each 29–30 (25–34) µm in diameter. Anal lobe well-developed, not very strong but considerably sclerotized, with 3 (2 or 3) enlarged conical setae with blunt apices on dorsal surface, ventrally with 1 or 2 (mostly 1) flagellate setae, each seta 49–54 (45–75) µm long; anal lobe with apical seta 160 (150–188) µm long. Dorsum. Derm membranous, smooth and without nodulations; anal lobes slightly sclerotized. Enlarged conical setae of 2 sizes: (i) short small setae, each approximately 11–20 (10–25) µm long, present on all dorsal surfaces except for body margin, but distinctly sparse on abdominal segments; (ii) longer conical setae with blunt apices, each 40–53 (30–61) µm long, present in transverse rows on cephalothorax and body margin. Margins of most abdominal segments each with 2 longer enlarged conical setae; submedial and medial areas of abdominal segment VII with 5 (4 or 5) shorter enlarged conical setae and abdominal segment VIII with 0 setae. Macrotubular ducts scattered on dorsal surface, each with outer ductule 10–11 (8–11) µm wide. Microtubular ducts simple, scattered throughout dorsum, each 5–10 µm long, elongate. Anal ring situated on dorsal margin, oval, slightly sclerotized, 50 (44–58) µm wide, 62 (56–68) µm long, partly with 2 rows of pores and 8 long setae; anal ring setae each 100–136 (88–137) µm long. Cauda absent. Venter. Derm membranous, smooth and without nodulation; anal lobes slightly sclerotized. Labium 3 segmented, joint length of 2 apical segments each about 110 (94–124) µm; basal segment with 2 pairs of setae, second segment with 2 (0–2) pairs of flagellate setae, apical segment with 3 (3–5) pairs of flagellate setae; stylet loop 2 to 3x longer than labium. Legs well developed, lengths of segments of prothoracic legs in µm: coxa, 56–61 (50–65); trochanter, 39–40 (34–50); femur, 130–132 (110–132); tibia, 88–89 (78–90); tarsus: 108–116 (101–116) and claw, 31–32 (24–32); length of tarsal digitules 50–57 (39–58); claw digitules 32–34 (18–35); length of segments of mesothoracic legs: coxa, 60–61 (53–65); trochanter, 39–40 (35–53); femur, 128–134 (102–134); tibia, 91–100 (80–100); tarsus, 111–119 (104–123) and claw, 30–32 (27–37); length of tarsal digitules 53–58 (49–58); claw digitules 38–39 (29–39); length of segments of metathoracic legs: coxa, 68–69 (57–72); trochanter, 42–43 (40–54); femur, 134–138 (105–138); tibia, 99–102 (84–102); tarsus, 120–123 (110–130) and claw, 34–37 (28–37); length of tarsal digitules 51–57 (51–60); claw digitules 29–39 (28–42); tarsal and claw digitules slightly knobbed; tarsal digitules longer than claw digitules; claw with a denticle; all coxae without spinulae. Tibia of each prothoracic leg with 5 (4–5) setae; tibia of each mesothoracic leg with 4 (3–4) setae; tibia of each metathoracic legs with 4 (2–5) setae. Multilocular disc pores each 5–6 (4–6) µm in diameter, mostly each with 5 loculi, present in relatively dense rows across abdominal segments, also a few on thoracic segments and head. Macrotubular ducts of 1 size, similar to dorsal macrotubular ducts but with outer ductules rather small and narrow; outer ductules each 6–7 µm wide, mostly present on submarginal to marginal areas. Microtubular ducts absent. A few cruciform pores present, mainly in submarginal areas of thoracic and anterior abdominal segments. A few enlarged conical setae with finely pointed apices present on marginal areas of thoracic segments and anterior abdominal segments; other setae flagellate, mainly present on medial to submedial areas of posterior body segments and on an area between antennae. Host plants. Rhododendron sp. (Ericaceae). Remarks. Acanthococcus torikurai sp. nov. is similar to Ac. glanduliferus Balachowsky 1933 in having two blunt enlarged setae on each margin of each abdominal segment, only a few enlarged setae on the dorsal abdominal segments, and the enlarged setae on the dorsum tending to get larger anteriorly. However, Ac. torikurai differs from Ac. glanduliferus as follows (contrasting character states in Ac. glanduliferus in parentheses): (i) in lacking small pores on hind legs (having small pores on hind legs); (ii) in lacking spinule on surface of meso- and metacoxae (having spinules on anterior surface of meso- and metacoxae); (iii) in lacking a cauda (having a cauda); (iv) in having different sized dorsomedial setae (dorsomedial setae are all of one size); (v) in lacking capitate ventral setae (having capitate ventral setae). Etymology. The new species is named after Mr Hidenori Torikura, the collector of the type series.

Genus Acanthococcus Signoret 1875 Type species: Acanthococcus aceris Signoret 1875 Genus diagnosis of adult female (adapted and slightly modified from Kozár et al. 2013). Ovisac ovoid, feltlike, white or gray, completely enclosing body of female. Adult female elongate-oval, narrowed posteriorly, with anal lobes conical and normally heavily sclerotized, sometimes strongly nodulose with sclerotized teeth on inner margin. Antennae 6, 7, or rarely 8 segmented; frontal lobes or tubercles usually present. Labium 3 segmented, with 2 well-developed distal segments; basal segment weakly developed and sclerotized with 2 pairs of hair-like setae; stylets short or very long, forming a double loop. Legs well developed, mid-coxa and hind coxa often with spinules on anterior surface, claw usually with a denticle; tarsal and claw digitules longer than claw, knobbed. Spiracles often each with a few associated multilocular disc pores; multilocular disc pores present on venter only, usually quinquelocular, but number of loculi varies between 3 and 9; oval disc pores (or cruciform pores) absent from dorsum, but often present on ventral surface of prosoma in a marginal band. Tubular ducts of 2 types, micro- and macrotubular ducts; microtubular ducts slender, scattered, or forming transverse rows or bands across dorsum, often associated with dorsal conical setae; macrotubular ducts often of 2 or 3 sizes, usually forming transverse rows or bands across body surfaces. Enlarged conical setae normally present at least on dorsal margin but often over entire dorsum, where they form transverse bands or rows; hair-like or flagellate setae present on venter only. Anal ring well developed, sclerotized with a partly double row of pores and usually 8 (rarely 6 or 10) anal ring setae each usually shorter than apical seta on anal lobe; each anal lobe with a long apical seta and usually with 3 short dorsal enlarged conical setae, seldom with more but at least with 2, ventral flagellate setae also present; cauda (median dorsal plate) usually well developed, but rarely absent. Remarks. The genus Acanthococcus is similar to Gossyparia Signoret 1875 in having enlarged conical setae present on most areas of the dorsum. It differs from Gossyparia in having dorsal macrotubular ducts present on most parts of dorsum, not restricted to the body margin, whereas Gossyparia has dorsal macrotubular ducts present mostly confined to the body margin., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2023, Two new species of Eriococcidae (Hemiptera: Coccomorpha) in Japan, pp. 387-395 in Zootaxa 5244 (4) on page 388, DOI: 10.11646/zootaxa.5244.4.5, http://zenodo.org/record/7663641, {"references":["Signoret, V. (1875) Essai sur les cochenilles ou gallinsectes (Homopteres - Coccides), 14 e partie. Annales de la Societe entomologique de France, Serie 5, 5, 15 - 40.","Kozar, F., Kaydan, M. B., Konczne Benedicty, Z. & Szita, E. (2013) Acanthococcidae and Related Families of the Palaearctic Region. Hungarian Academy of Sciences, Budapest, 680 pp."]}

Genus Anophococcus Balachowsky 1954 Type species: Eriococcus inermis Green 1915. Genus diagnosis of adult female (adapted and slightly modified from Kozár et al. 2013). Ovisac ovoid, felt-like, white or gray, completely enclosing body of female. Adult female elongate oval, narrowed posteriorly, with anal lobes conical, slightly sclerotized. Antennae usually 6 or 7 segmented; frontal lobes usually absent; frontal tubercles usually present, rarely absent. Labium 3 segmented, basal segment weakly developed and sclerotized, with 2 pairs of flagellate setae; apical setae of labium each approximately half length of the subapical setae, or less. Stylets usually short, about as long as labium. Legs well-developed: mesocoxae and metacoxae often with spinules on anterior surfaces; translucent sensory pores present or absent; claws usually each with a denticle; tarsal and claw digitules mostly longer than claw, usually knobbed but rarely spine-like. Spiracles often each with a few associated multilocular disc pores; multilocular disc pores present on venter only, usually each with 5 loculi, but number of loculi varies between 3 to 9; cruciform pores absent from dorsum, but often present on venter of prosoma in a marginal band, not grouped. Tubular ducts of 2 types, micro- and macrotubular ducts; microtubular ducts usually short, scattered, or forming transverse rows or bands on dorsum, often among dorsal conical setae; macrotubular ducts often of 2 or 3 sizes, usually forming transverse rows or bands on both body surfaces. Enlarged conical setae normally present only on margins, sometimes only on anal lobes and on margin of head; dorsal setae usually minute but sometimes setae in some areas more developed. Flagellate setae present on venter only. Anal ring welldeveloped, sclerotized with a single or only partly double row of pores and with 8 (rarely 6) anal ring setae, each as long as, or shorter than apical seta on anal lobes; each anal lobe with a long apical seta and usually 3 short dorsal conical setae, seldom with 4 setae. Cauda usually absent, but rarely present. Remarks. The genus Anophococcus is similar to the genera Acanthococcus Signoret 1875, Gossyparia Signoret 1875, Rhizococcus Signoret 1875 and Uhleria Cooke 1881, in lacking cruciform pores or discoidal pores on the dorsum. However, Anophococcus differs from the four other genera in having (contrasting character states in Acanthococcus, Gossyparia, Rhizococcus and Uhleria in parentheses): dorsal enlarged conical setae confined to body margin or restricted to anal lobe (dorsal enlarged conical setae present on most or all of anterior half (cephalothoracic segments)). This morphological character state can sometimes be challenging to discern and may be considered an artificial criterion. Some Anophococcus species, such as An. pseudinsignis (Green 1921) and An. sanguinairensis (Goux 1993), possess well-developed and long dorsal enlarged conical setae not only on the margin but also on the head apex. Additionally, in Acanthococcus torikurai sp. nov. and Rhizococcus coccineus (Cockerell 1894), dorsal conical setae are greatly reduced in the medial and submedial areas of the dorsal abdominal segments, and the setae are obviously smaller and shorter than the marginal dorsal enlarged conical setae. The new species we propose below also has relatively well-developed dorsal enlarged conical setae on the head apex. There may be some uncertainty as to whether consistent morphological differences exist between the species of the genera Acanthococcus and Rhizococcus and the new species. However, since the new species has short microtubular ducts and shares this morphological character state with other species of the genus Anophococcus, it is tentatively placed in the genus Anophococcus in this study., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2023, Two new species of Eriococcidae (Hemiptera: Coccomorpha) in Japan, pp. 387-395 in Zootaxa 5244 (4) on page 391, DOI: 10.11646/zootaxa.5244.4.5, http://zenodo.org/record/7663641, {"references":["Balachowsky, A. S. (1954) Sur une nouvelle espece d'Eriococcini de la foret de Fontainebleau avec creation d'un genre nouveau. [Hom. Coccoidea.]. Bulletin de la Societe entomologique de France, 59, 61 - 64. https: // doi. org / 10.3406 / bsef. 1954.18703","Green, E. E. (1915) Observations on British Coccidae in 1914, with descriptions of new species. Entomologist's Monthly Magazine, Series 3, 51, 175 - 185. https: // doi. org / 10.5962 / bhl. part. 7788","Kozar, F., Kaydan, M. B., Konczne Benedicty, Z. & Szita, E. (2013) Acanthococcidae and Related Families of the Palaearctic Region. Hungarian Academy of Sciences, Budapest, 680 pp.","Signoret, V. (1875) Essai sur les cochenilles ou gallinsectes (Homopteres - Coccides), 14 e partie. Annales de la Societe entomologique de France, Serie 5, 5, 15 - 40.","Cooke, M. (1881) A treatise on the insects injurious to fruit and fruit trees of the state of California and remedies recommended for their extermination. State Office Sacramento, Sacramento, California, 72 pp. https: // doi. org / 10.5962 / bhl. title. 37277","Green, E. E. (1921) Observations on British Coccidae with descriptions of new species. No. VI. Entomologist's Monthly Magazine, Series 3, 57, 146 - 152, 189 - 200. https: // doi. org / 10.5962 / bhl. part. 22330","Goux, L. (1993) Description de trois especes nouvelles d' Eriococcus appartenant a la faune Francaise (Homoptera, Coccoidea, Eriococcidae). Bulletin de la Societe Linneenne de Provence, 44, 65 - 69.","Cockerell, T. D. A. (1894) Descriptions of new Coccidae. Entomological News, 5, 203 - 204."]}

Key to adult females of the family Eriococcidae found in Japan (adapted and modified from Kawai 1980 and Kozár et al. 2013) 1(0) Dorsum without macrotubular ducts, but with large invaginated 5-locular pores. Labium small, reduced, indistinctly two segmented, with basal segment absent. Metathoracic legs greatly reduced, each represented by a ball-shaped sclerotization............................................................................ Kuwanina parva (Maskell 1897) - Dorsum with macrotubular ducts, but without large invaginated 5-locular pores. Labium large, well developed, with basal segment present. Metathoracic legs well developed.......................................................... 2 2(1) Basal segments of labium with 0 or 1 pair of setae. Antenna 5 segmented. Dorsum with conical setae broad............................................................................... Asiacornococcus japonicus (Kuwana 1902) - Basal segments of labium with 2 pairs of setae. Antenna with more than 5 segments. Dorsal conical setae relatively slender, not broad.............................................................................................. 3 3(2) Dorsal macrotubular ducts present on margin only........................................................... 4 - Dorsal macrotubular ducts present over entire dorsum........................................................ 5 4(3) Dorsal setae numerous....................................................... Gossyparia spuria (Modéer 1778) - Dorsal setae few, scarcely present........................................ Acalyptococcus graminis (Maskell 1897) 5(3) Enlarged conical setae quite small in medial and mediolateral areas of dorsum (each less than half as long as a marginal seta), especially on abdomen................................................. Anophococcus koguchii Tanaka sp. nov. - Enlarged conical setae medium-sized in medial and mediolateral areas of dorsum (each about half as long as a marginal seta)................................................................................................... 6 6(5) Microtubular ducts short. Found mostly on herbaceous plants except bamboo..................................... 7 - Microtubular ducts long. Found mostly on woody plants and bamboo.......................................... 11 7(6) Body elongate oval, with body length about 1.5x body width, always less than 2x body width. Found on Asteraceae, Cactaceae, and Fabaceae........................................................................................ 8 - Body extremely elongate oval, with body length about 2.5x body width. Found on Poaceae......................... 10 8(7) Metathoracic coxa much larger than prothoracic or mesothoracic coxa, wider than length of metathoracic tibia...................................................................................... Rhizococcus sojae Kuwana 1917 - Metathoracic coxa about same size as prothoracic or mesothoracic coxa, narrower than length of metathoracic tibia....... 9 9(8) Conical setae of medial area of posterior abdominal segments quite small................. R. coccineus (Cockerell 1894) - Conical setae of medial area of posterior abdominal segments well developed and as large as marginal enlarged conical setae............................................................................... R. henmii (Kuwana 1931) 10(7) Dorsal surface of each anal lobe with 4 enlarged conical setae........................... R. miscanthi Takahashi 1957 - Dorsal surface of each anal lobe with 3 enlarged conical setae........................... R. festucae (Lindinger 1932) 11(6) Dorsum with enlarged conical setae blunt or truncate....................................................... 12 - Dorsum with enlarged conical setae sharply pointed........................................................ 14 12(11) Dorsum of abdominal segments with conical setae small and scarce............. Acanthococcus torikurai Tanaka sp. nov. - Dorsum of abdominal segments with conical setae not small or scarce.......................................... 13 13(12) Thoracic submargin of venter with macrotubular ducts................................. Ac. tokaedae (Kuwana 1932) - Thoracic submargin of venter without macrotubular ducts...................... Ac. chabohiba (Kuwana & Nitobe 1918) 14(11) Inner margin of anal lobe with some granule-like sclerotizations. Found on bamboo............................... 15 - Inner margin of anal lobe without granule-like sclerotizations. Found on woody plants............................. 16 15(14) Dorsum of abdominal segment VIII with fewer than 10 conical setae........................ Ac. onukii (Kuwana 1902) - Dorsum of abdominal segment VIII with more than 20 conical setae........................ Ac. azumae (Kanda 1933) 16(14) Anal lobe with most dorsal setae all about same length................................ Ac. abeliceae (Kuwana 1927) - Anal lobe with dorsal setae on basal inner margin smaller than other 2 setae........... Ac. lagerstroemiae (Kuwana 1907), Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2023, Two new species of Eriococcidae (Hemiptera: Coccomorpha) in Japan, pp. 387-395 in Zootaxa 5244 (4) on pages 393-394, DOI: 10.11646/zootaxa.5244.4.5, http://zenodo.org/record/7663641, {"references":["Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Kozar, F., Kaydan, M. B., Konczne Benedicty, Z. & Szita, E. (2013) Acanthococcidae and Related Families of the Palaearctic Region. Hungarian Academy of Sciences, Budapest, 680 pp.","Maskell, W. M. (1897) On a collection of Coccidae, principally from China and Japan. Entomologist's Monthly Magazine, 33, 239 - 244. https: // doi. org / 10.5962 / bhl. part. 18316","Kuwana, S. I. (1902) Coccidae (scale insects) of Japan. Proceedings of the California Academy of Sciences, 3, 43 - 98.","Modeer, A. (1778) Om fastflyet. Coccus. Handlingar Gotheborgska Wetenskaps och Witterhets Samhallets. Wetenskaps Afdelingen. Gotheborg, Series 1, 1, 11 - 50.","Kuwana, S. I. (1917) [About the genus Eriococcus in Japan]. Konchuu Sekai, 21 (236), 1 - 4. [in Japanese]","Cockerell, T. D. A. (1894) Descriptions of new Coccidae. Entomological News, 5, 203 - 204.","Kuwana, S. I. (1931) Scale insects of Amami-Oshima, with descriptions of four new species. Dobutsugaku Zasshi (Journal of the Zoological Society of Japan). Tokyo, 43, 163 - 171.","Takahashi, R. (1957) A new species of Eriococcus from Japan (Coccoidea, Homoptera). Akitu, Transactions of the Kyoto Entomological Society, 6, 6 - 7.","Lindinger, L. (1932) Beitrage zur Kenntnis der Schildlause. Konowia, 11, 177 - 205.","Kuwana, S. I. (1932) Description of two new Coccidae from Japan. Kontyu, 6, 143 - 148. Kuwana, S. I. & Nitobe, I. (1918) On a new species of Eriococcus from Japan. Insect World, 22, 400 - 403.","Kanda, S. (1933) A new species of the genus Eriococcus. Insect World, 37, 151 - 153.","Kuwana, S. I. (1927) A new Eriococcus from Japan. Dobutsugaku Zasshi (Journal of the Zoological Society of Japan). Tokyo, 39, 109 - 113.","Kuwana, S. I. (1907) Coccidae of Japan, I. A synoptical list of Coccidae of Japan with descriptions of thirteen new species. Bulletin of the Imperial Central Agricultural Experiment Station, Japan, 1, 177 - 212."]}

Genus Spilococcus Ferris 1950 Spilococcus Ferris 1950: 219. Type species: Dactylopius gutierreziae Cockerell 1896. Genus diagnosis (modified from McKenzie 1967). Body of adult female elongate oval to broadly oval; anal lobe developed, each lobe without anal lobe bar and bar seta, usually with 2 conical setae, 1 or more slender auxiliary setae, and a concentration of trilocular pores. Other cerarii usually numbering 6–17 pairs, each containing 2 conical setae and a slight concentration of trilocular pores. Antennae each usually with 8 segments, slender. Legs well developed, translucent pores often present on hind coxae and hind tibiae. Tarsal digitules knobbed. Circulus present or absent. Anterior and posterior ostioles usually present. Anal ring normal, usually bearing 6 setae. Body setae often flagellate, sometimes conical or spiniform. Trilocular pores normal. Multilocular disc pores present, usually confined to venter. Oral rim tubular ducts present at least on dorsum. Oral collar tubular ducts present on venter at least. Remarks. The genus Spilococcus is similar to Paracoccus Ezzat & McConnell 1956 in having oral rim tubular ducts and a small number of cerarii (Spilococcus should be transferred to Paracoccus (see “Remarks” under Spilococcus pacificus). Further morphological and molecular phylogenetic studies including these two genera are greatly needed. The separation of Crisicoccus from Spilococcus is discussed in “Remarks” under C. seruratus above., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on page 567, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Ferris, G. F. (1950) Atlas of the Scale Insects of North America. Series 5. Vol. 5. The Pseudococcidae (Part I). Stanford University Press, Palo Alto, California, 278 pp.","Cockerell, T. D. A. (1896) New species of insects taken on a trip from the Mesilla Valley to the Sacramento Mts., New Mexico. Journal of the New York Entomological Society, 4, 201 - 207.","McKenzie, H. L. (1967) Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley, 526 pp. https: // doi. org / 10.1525 / 9780520338227","Ezzat, Y. M. & McConnell, H. S. (1956) A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A-e 84, 1 - 108."]}

Key to separate the genera Paracoccus, Spilococcus and other related genera with oral rim tubular ducts (adapted and modified from Kaydan & Szita 2017) 1(0) Oral rim tubular ducts present, each with well-developed rim.................................................. 2 - Oral rim tubular ducts absent entirely.................................................... rest of Pseudococcinae 2(1) Cerarii entirely absent............................................. Humococcus Ferris and Bromusicoccus Kaydan - At least anal lobe cerarii present......................................................................... 3 3(2) Multilocular disc pores absent from venter except occasionally for 1 or 2 around vulva.............. Distichlicoccus Ferris - Multilocular disc pores present on venter at least around vulva................................................. 4 4(3) Some abdominal cerarii each with 3 or more conical setae................................................................................................... Moystonia Williams and Brasiliputo Williams & Granara de Willink - All abdominal cerarii each with 2 conical setae.............................................................. 5 5(4) Cerarii numbering no more than 6 pairs, on abdomen only............................................................ Maconellicoccus Ezzat, Atrococcus Goux, Colombiacoccus Williams & Granara de Willink, and Vryburgia De Lotto. - Cerarii numbering 9–18 pairs, not confined to abdomen....................................................... 6 6(5) Legs usually long and slender, extending well beyond margins of body; claws noticeably long, pointed and slender, usually straight; rims of oral rim tubular ducts, although well developed, fairly flat in profile.................. Leptococcus Reyne - Legs rarely extending beyond margins of body; claws stout and curved; rims of oral rim tubular ducts elevated above derm when viewed in profile................................................................................. 7 7(6) Venter of each anal lobe with an anal lobe bar, present either forwards from apical seta or from bar seta only........................................................................................ Paracoccus Ezzat & McConnell - Venter of each anal lobe without anal lobe bar, although differently shaped sclerotized area may be present.............. 8 8(7) Venter of each anal lobe with triangular-quadrate sclerotized area occupying much of lobe; auxiliary setae present at least in anal lobe cerarii and penultimate cerarii (C 17 and C 18)...................................... Pseudococcus Westwood - Venter of each anal lobe without any sclerotized area; auxiliary setae present in anal lobe cerarii only..... Spilococcus Ferris

Crisicoccus ezzati sp. nov. [Japanese common name: Nashi-kona-kaigaramushi] Crisicoccus matsumotoi (Shiraiwa 1935); Ezzat & McConnell 1956: 25; Williams 2004: 144 (misidentifications). Material examined. Holototype, here designated: Pseudococcus / Crisicoccus / matsumotoi / (Shir.) / On Pears / Japan: at Seattle / Berryhill. Scott. Collr’s / Sept. 12. 1940 / Seattle 9045. 1 adult female mounted singly on a slide (USNM). Description based on the holotype only. Appearance in life. Not seen. Slide-mounted adult female (Fig. 3). Body elongate oval, 2.0 mm long and 1.0 mm wide; derm membranous; segmentation recognizable, but not well developed. Anal lobes well developed, ventral surface with rather faint anal lobe bar and a long apical seta, 226–254 µm long. Antenna 399–410 µm long, with 8 segments and many flagellate setae; subapical segment with 1 fleshy seta and apical segment with 3 fleshy setae. Eyespot present on margin, not associated with discoidal pores. Legs well developed, with many flagellate setae; hind leg measurements (in µm): trochanter + femur 304–307 long; hind tibia + tarsus 304–312; claw 34–35, without a denticle. Ratio of lengths of hind tibia + tarsus to trochanter + femur about 1: 0.99–1.03; ratio of lengths of hind tibia to tarsus 1: 2.12–2.23. Paired setose tarsal digitules present, subequal in length to the minutely knobbed claw digitules. Hind coxa with translucent pores on both surfaces; hind tibia with translucent pores on posterior surface; hind trochanter, femur and tarsus without translucent pores. Labium 218 µm long, shorter than clypeolabral shield. Circulus usually quadrate, located between abdominal segments III and IV, 76 µm long and 109 µm wide. Ostioles present, each with inner edges of lips weakly sclerotized; anterior ostioles each with a total for both lips of 23 trilocular pores and 2 setae; each posterior ostiole with a total for both lips of 25–28 trilocular pores and 3–5 setae. Anal ring 95 µm wide, bearing 6 setae, each seta 108–138 µm long. Cerarii numbering 15–16 pairs. Anal lobe cerarii (C 18) usually each containing 2 conical cerarian setae, each seta 20–22 µm long and about 5–6 µm wide at base, 6–7 auxiliary setae and a concentration of trilocular pores. Penultimate cerarii (C 17) each containing 2 conical setae and a few auxiliary setae and trilocular pores. Cerarii situated further forward generally each with 2–4 conical setae, usually with flagellate tips, and trilocular pores. Dorsum. Setae flagellate, each 14–48 µm long, distributed segmentally; longest setae present on head. Trilocular pores each 3–4 µm wide, evenly distributed. Oral rim tubular ducts and oral collar tubular ducts absent. Discoidal pores each about 2 µm wide, sparsely distributed. Venter. Setae relatively long and flagellate, each 18–89 µm long; setae on medial area of posterior abdominal segments longest. Multilocular disc pores, each 6–8 µm wide, present in medial areas of abdominal segments IV‒IX, arranged in 1 (or rarely 2) rows on each posterior area of abdominal segments IV–VII but arranged randomly on abdominal segments VIII and IX. Trilocular pores, each same size as those on dorsum, evenly distributed. Oral collar tubular ducts all of 1 size, each narrower than a trilocular pore, about 2–3 µm wide, present on medial areas of abdominal segments IV–VII and submarginal to marginal areas of abdominal segments II–VIII, usually forming transverse bands across segments; additionally, a few ducts present on submarginal to marginal areas of thoracic segments. Discoidal pores, same size as those on dorsum, sparsely present. Host plant in Japan. Rosaceae: Pyrus pyrifolia var. culta. Remarks. Crisicoccus ezzati sp. nov. was first reported as C. matsumotoi by Ezzat & McConnell (1956), based on specimens from Japan intercepted at a US plant quarantine station and three specimens collected in Fukuoka prefecture, Japan. Some specimens collected later from India and Philippines were reported by Williams (2004). In this study, we examined a specimen identified by Ezzat & McConnell (1956) and found that the species is quite different from that currently considered to be C. matumotoi (= Crisicoccus seruratus) in Japan. Crisicoccus ezzati has (contrasting characteristics of C. seruratus are stated in parentheses): (i) cerarii numbering 15–16 pairs (no more than eight pairs of cerarii); and (ii) only one size of oral collar tubular ducts on the venter (two sizes of oral collar tubular ducts on the venter). Judging the validity of this species being labelled “ C. matsumotoi ” is difficult because the type specimens for C. matsumotoi are lost (García Morales et al. 2016). However, in this study, we followed Dr Shozo Kawai, Professor Emeritus of Tokyo University of Agriculture, who studied Japanese mealybugs taxonomically for a long time and treated the species currently recognized as C. matsumotoi in Japan as the true C. matsumotoi (= C. seruratus). Therefore, the species described by Ezzat & McConnell (1956) and Williams (2004) lacks a valid name and is here described as a new species. Crisicoccus ezzati bears some similarities to C. orchidiradicis (Takahashi 1951) from Malaysia in having a large number of cerarii (≥ 15 pairs) and anterior cerarii with relatively narrow, elongated cerarian setae. However, it differs by the following morphological characteristics (contrasting characteristics of C. orchidiradicis are given in parentheses): (i) 15–16 pairs of cerarii (17 pairs); (ii) translucent pores present on hind coxae (without translucent pores on hind coxae), and (iii) anterior cerarii lacking auxiliary setae (all cerarii containing auxiliary setae). Etymology. The new species is named after Dr Yehia Mahmoud Ezzat, who was one of the first to report on it., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on pages 560-562, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Shiraiwa, H. (1935) Studies on mealybugs infesting pear in Japan. Kontyu, 9, 63 - 75.","Ezzat, Y. M. & McConnell, H. S. (1956) A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A-e 84, 1 - 108.","Williams, D. J. (2004) Mealybugs of southern Asia. The Natural History Museum, London and Southdene SDN. BHD., Kuala Lumpur, 896 pp.","Takahashi, R. (1951) Some mealy bugs (Pseudococcidae, Homoptera) from the Malay Peninsula. Indian Journal of Entomology, 12 (1950 - 1), 1 - 22."]}

Crisicoccus pini (Kuwana 1902) [Japanese common name: Matsu-kona-kaigaramushi] Dactylopius pini Kuwana 1902: 54. Pseudococcus pini (Kuwana 1902); Fernald 1903: 107 (change of combination). Crisicoccus pini (Kuwana 1902); Ferris 1950: 46 (change of combination); McKenzie 1967: 127; Kawai 1972: 6; Kawai 1980: 107; Tang 1992: 356; Kosztarab 1996: 94; Kawai 2003: 929; Kwon et al. 2003: 404; Danzig & Gavrilov-Zimin 2015: 202. Material examined. Japan: Shizuoka Prefecture, Shimizu, Okitsu, on Pinus sp., 2.vi.1972, coll. S. Kawai, 6 adult females mounted on 2 slides (KTUA); Fukuoka Prefecture, Fukuoka, Nishi-ku, Ikinomatsubara, on Pinus thunbergii, 9.v.2021, coll. H. Tanaka, 14 adult females mounted singly (7 ELKU, 7 EUMJ). Updated description Appearance in life (Fig. 5). Adult female 2‒4 mm long, light orange, covered with white powdery wax. Wax projections from body margin short and indistinct on cephalothorax, and slightly longer on a few segments on posterior part of body. Slide-mounted adult female (Fig. 6) (n = 20). Body elongate oval, 2.6–4.0 mm long and 1.2–2.0 mm wide; derm membranous; segmentation recognizable, but not well developed. Anal lobes well developed, ventral surface with slightly faint, narrow anal lobe bar and a long apical seta, 130–228 µm long. Antenna 328–451 µm long, with 8 segments and many flagellate setae; subapical segment with 1 fleshy seta and apical segment with 3 fleshy setae. Eyespots present on margin, not associated with discoidal pores. Legs well developed, with many flagellate setae; hind leg measurements (in µm): trochanter + femur 243–348 long; hind tibia + tarsus 255–379; claw 24–34, without a denticle. Ratio of lengths of hind tibia + tarsus to trochanter + femur about 1: 0.91–1.15; ratio of lengths of hind tibia to tarsus 1: 1.90–2.87. Paired setose tarsal digitules present, subequal in length to the minutely knobbed claw digitules. Hind coxa with translucent pores on both surfaces; hind tibia occasionally with translucent pores on posterior surface; hind trochanter, femur and tarsus without translucent pores. Labium 98–165 µm long, shorter than clypeolabral shield. Circulus lacking. Ostioles present, each with inner edges of lips weakly sclerotized; anterior ostioles each with a total for both lips of 12–27 trilocular pores and 2–8 setae; each posterior ostiole with a total for both lips of 13–30 trilocular pores and 2–7 setae. Anal ring 68–88 µm wide, bearing 6 setae, each seta 98–174 µm long. Cerarii numbering 4–7 pairs, all present on abdominal segments.Anal lobe cerarii (C 18) mostly each containing 2 cerarian setae, each seta 14–40 µm long and about 4–8 µm wide at base, 1–6 auxiliary setae and a concentration of trilocular pores. Penultimate cerarii (C 17) each containing 2 conical setae, no auxiliary setae and a few trilocular pores. Cerarii situated further forward generally each with 2 conical setae with a few trilocular pores. Dorsum. Setae relatively short and flagellate, each 6–39 µm long, usually distributed segmentally; longest setae present on head. Trilocular pores each 3–4 µm wide, evenly distributed. Oral rim tubular ducts absent. Oral collar tubular ducts present or absent, if present, each 2–3 µm wide, located on marginal to submarginal area of posterior abdominal segments. Discoidal pores each about 1–2 µm wide, sparsely distributed. Venter. Setae relatively long and flagellate, each 7–122 µm long; setae on head longest. Multilocular disc pores, each 6–10 µm wide, present in medial areas of abdominal segments IV‒IX, but occasionally absent from a few anterior segments, arranged in 1–2 rows on posterior area of abdominal segment VII; 0–1 row on anterior area of abdominal segment VII; in 1 row on each posterior area of abdominal segments IV‒ VI but arranged randomly on abdominal segments VIII and IX. Trilocular pores, each same size as those on dorsum, evenly distributed. Oral collar tubular ducts all 1 size, each about 2–3 µm wide, present on medial areas of abdominal segments III–VII and usually forming transverse bands across segments, also present on submarginal to marginal areas of posterior abdominal segments. Discoidal pores, same size as those on dorsum, sparsely present. Host plants in Japan. Pinaceae: Pinus densiflora, P. parviflora and P. thunbergii (Kawai 1972; 1980; 2003). Remarks. Crisicoccus pini is similar to C. azaleae in having a small number of cerarii (C. azaleae above., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on pages 564-567, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Kuwana, S. I. (1902) Coccidae (scale insects) of Japan. Proceedings of the California Academy of Sciences, 3, 43 - 98.","Fernald, M. E. (1903) A catalogue of the Coccidae of the world. Bulletin of the Hatch Experiment Station of the Massachusetts Agricultural College, 88, 1 - 360. https: // doi. org / 10.5962 / bhl. title. 8533","Ferris, G. F. (1950) Atlas of the Scale Insects of North America. Series 5. Vol. 5. The Pseudococcidae (Part I). Stanford University Press, Palo Alto, California, 278 pp.","McKenzie, H. L. (1967) Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley, 526 pp. https: // doi. org / 10.1525 / 9780520338227","Kawai, S. (1972) Diagnostic notes and biology of the coccid species occurring on cultivated or wild trees and shrubs in Japan (Homoptera: Coccoidea). Bulletin of the Tokyo-To Agricultural Experiment Station, 6, 1 - 54. [in Japanese]","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Tang, F. T. (1992) [The Pseudococcidae of China.] Chinese Agricultural Science Technology Press Beijing, 768 pp. [in Chinese with English abstract]","Kosztarab, M. P. (1996) Scale insects of Northeastern North America. Identification, biology, and distribution. Virginia Museum of Natural History, Martinsburg, Virginia, 650 pp.","Kawai, S. (2003) Kaigaramushi [Coccoidea]. In: Umeya, K. & Okada, T. (Eds.), Agricultural insect Pests in Japan. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 1 - 1203. [in Japanese]","Danzig, E. M. & Gavrilov-Zimin, I. A. (2015) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 2: Subfamily Pseudococcinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 619 pp."]}

Crisicoccus seruratus (Kanda 1933) [Japanese common name: Matsumoto-kona-kaigaramushi] Pseudococcus seruratus Kanda 1933: 133. Pseudococcus matsumotoi Shiraiwa 1935: 66. syn. nov. Pseudococcus astericola Shinji 1936: 49 (synonymy by Kanda 1941: 24). Crisicoccus seruratus (Kanda 1933); Paik 1978: 186 (change of combination); Kawai 2003: 314. Crisicoccus matsumotoi (Shiraiwa 1935); Kawai 1980: 107 (change of combination). Material examined. Neotype, here designated: JAPAN / Tokyo, Tachikawa, / Fujimi-cho, / on Zelkova serrata, / 8.v.1962, / coll. S. Kawai; an adult female mounted with other 9 adult females on a slide (KTUA). The individual at the left end of the middle row of specimens was selected as the neotype and is clearly indicated on the slide with a black ink dot (Fig. 1). Other material. JAPAN: Tokyo, Chuo-ku, Hamarikyuteien, on Mallotus japonicus, 15. v.1972, coll. S. Kawai, 6 adult females mounted on 2 slides (KTUA); Shizuoka prefecture, Shimizu, Okitsu, on Diospyros kaki, 8.vii.1976, coll. S. Kawai, an adult female mounted singly (KTUA); Shimane prefecture, Izumo, on Vitis sp., 25.v.2001, coll. S. Narai, an adult female mounted singly (KTUA); Ibaraki prefecture, Kasama, Ago, on Pyrus pyrifolia var. culta, 27.iv.2022, coll. T. Tsunoda, 6 adult females mounted singly (3 ELKU, 3 EUMJ). Updated description Appearance in life. Adult female 3‒4 mm long, dark purple covered with a white powdery wax. Projections of the wax secretion from body margin not so developed and limited to a few segments of posterior part of body. The body contents of this species turn blue black to dark green in 10% potassium hydroxide solution (Kawai 1980, translated by HT). Slide-mounted adult female (Fig. 2) (n = 15). Body elongate oval, 3.6 (1.9–4.5) mm long and 2.2 (1.2–3.0) mm wide; derm membranous; segmentation recognizable, but not well developed. Anal lobes well developed, each with a long apical seta, 188–242 (145–270) µm long. Each lobe with a narrow and slightly faint anal lobe bar on ventral surface, but bar sometimes difficult to see in some specimens. Antenna 368–369 (368–500) µm long, with 8 segments and many flagellate setae; subapical segment with 1 fleshy seta and apical segment with 3 fleshy setae. Eyespot present on margin, not associated with discoidal pores. Legs well developed, with many flagellate setae; hind leg measurements (in µm): trochanter + femur 269–270 (269–394) long; hind tibia + tarsus 281–286 (281–377); claw 39–42 (34–45), without a denticle. Ratio of lengths of hind tibia + tarsus to trochanter + femur about 1: 1.04–1.06 (1: 0.89–1.06); ratio of lengths of hind tibia to tarsus 1: 1.78–1.83 (1: 1.78–2.60). Paired setose tarsal digitules present, subequal in length to the minutely knobbed claw digitules. Hind coxa with translucent pores on both surfaces; hind tibia usually with translucent pores on posterior surface; hind trochanter, femur and tarsus without translucent pores. Labium 170 (127–185) µm long, mostly shorter than clypeolabral shield. Circulus oval to quadrate, located between abdominal segments III and IV and divided by an intersegmental line, 70 (50–170) µm long and 90 (48–165) µm wide. Ostioles present, each with inner edges of lips weakly sclerotized; anterior ostioles each with a total for both lips of 22–27 (11–59) trilocular pores and 5 (2–7) setae; each posterior ostiole with a total for both lips of 24–27 (19–62) trilocular pores and 5–8 (2–8) setae. Anal ring 94 (90–113) µm wide, bearing 6 (6–8) setae, each seta 112–145 (60–170) µm long. Cerarii usually numbering 4 or 5 (4–8) pairs, mostly present on posterior abdominal segments, rarely present also on thoracic segments. Anal lobe cerarii (C 18) each containing 2 conical cerarian setae, each seta 17–20 (9–23) µm long and about 7 (5–8) µm wide at base, 4–5 (4–8) auxiliary setae and a concentration of trilocular pores. Penultimate cerarii (C 17) each containing 2 conical setae, 3 (0–8) auxiliary setae and a few trilocular pores. Cerarii situated further forward generally each with 2 conical setae with a few trilocular pores and auxiliary setae. Dorsum. Setae flagellate, each 17–70 µm long, distributed usually segmentally; longest setae present on head. Trilocular pores each 3–4 µm wide, evenly distributed. Oral rim tubular ducts and oral collar tubular ducts absent. Discoidal pores each about 2–3 (1–4) µm wide, sparsely distributed. Venter. Setae relatively long and flagellate, each 32–90 (19–168) µm long; setae on head longest. Multilocular disc pores, each 6–8 (6–9) µm wide, present in medial areas of abdominal segments IV‒IX, arranged in single row on each posterior area of abdominal segments IV‒ V, in 2 or 3 rows on each posterior area of abdominal segments VI –VII, and in 1–2 rows on anterior areas of abdominal segments VI –VII but arranged randomly on abdominal segments VIII and IX. Trilocular pores, each same size as those on dorsum, evenly distributed. Oral collar tubular ducts of 2 sizes present: (i) large-type ducts, each about 3–4 µm in diameter, mostly wider than a trilocular pore, present on marginal to submarginal areas of all abdominal and thoracic segments and on medial areas of abdominal segments IV– VII, also forming transverse bands across abdominal segments IV–VII; and (ii) small-type ducts, each about 2–3 μm in diameter, mostly smaller than a trilocular pore, present on medial areas of abdominal segments IV–VII intermixed with large-type ducts, forming transverse bands across abdominal segments IV–VII. Discoidal pores, same size as those on dorsum, sparsely present. Host plants in Japan. Buxaceae: Buxus microphylla (Kawai 1980); Cannabaceae: Celtis sinensis (Kawai 1980); Ebenaceae: Diospyros kaki (Kawai 1980; Kawai 2003); Euphorbiaceae: Mallotus japonicus (Kawai 1980); Fabaceae: Wisteria floribunda (Kawai 2003); Juglandaceae: Juglans mandshurica (Kawai 1980); Moraceae: Ficus carica (Kawai 1980; Kawai 2003); Platanaceae: Platanus orientalis (Kawai 1980); Rosaceae: Chaenomeles speciosa (Kawai 1980), Photinia glabra (Kawai 2003), Pyrus pyrifolia var. culta (Kawai 1980; Kawai 2003); Sapindaceae: Acer palmatum (Kawai 1980), Acer spp. (Kawai 2003); Ulmaceae: Zelkova serrata (Kawai 1980); Vitaceae: Vitis spp. (Kawai 2003). Remarks. The original description of C. seruratus by Kanda (1933) was based mostly on highly variable morphological characteristics (including proportions of antennal segment lengths) that have little taxonomic value. Most of Kanda’s specimens are deposited in Osaka Museum of Natural History, Japan (OMNH); however, a lengthy search was made for type specimens of this species in almost all of the Coccomorpha collections in Japan (including Osaka Museum of Natural History) but none could be found; we therefore conclude that all the type specimens of C. seruratus described by Kanda have been lost. The designation of a neotype of this species was necessary because the original description is not informative, complicated taxonomic problems are associated with the species, and the type material has been lost. One of the specimens used in this redescription is designated the neotype (above) for taxonomic stability. The neotype specimen was obtained from a tree of Zelkova serrata in the Kanto region of Japan, which is the same area where the primary type series for C. seruratus were collected. The morphology of this type specimen agrees well with the original description. Currently, in Japan C. seruratus is recognized as C. matsumotoi. Its morphology differs significantly from “ C. matsumotoi ” reported by Ezzat & McConnell (1956) and Williams (2004), by having the following morphological characteristics (contrasting characteristics of the material incorrectly identified as “ C. matsumotoi ” are stated in parentheses): (i) a small number of cerarii, numbering fewer than 8 pairs (cerarii numbering 15–16 pairs); and (ii) oral collar tubular ducts of two different sizes present on venter (venter with only one size of oral collar tubular ducts). Therefore, in this study, we describe “ C. matsumotoi ” reported by Ezzat & McConnell (1956) and Williams (2004) as a distinct, new species, Crisicoccus ezzati sp. nov. (see below). Crisicoccus seruratus has been collected on the roots of Zelkova serrata (= Abelicea serrata), which is the host plant of Kanda’s original collection, but he used the name Abelicea serrata and he misspelt the scientific name of the host as “ Obelicea serurata ”. Kawai's (1980) view that C. matsumotoi and C. seruratus are the same species seems to be correct; however, he incorrectly designated C. seruratus (the older name) as a junior synonym of C. matsumotoi (García Morales et al. 2016). Above, we formally synonymize Pseudococcus matsumotoi Siraiwa 1935 as a junior synonym of Pseudococcus seruratus Kanda. We were unable to examine any types of Pseudococcus astericola Shinji 1936; however, Kanda (1941) synonymized this species under Pseudococcus seruratus Kanda and we follow his synonymy. In Korea, Paik (1978) and Kwon et al. (2003) regarded C. seruratus and “ C. matsumotoi ” as distinct species; however, the species they recognized as “ C. matsumotoi ” in Korea was a misidentification of Spilococcus pacificus (see below). Crisicoccus species do not have oral rim tubular ducts on the body surface, whereas Spilococcus species have at least a few so the genera can be easily distinguished from each other. Crisicoccus seruratus shows similarities to C. melaleucae Williams 1985 (described from Queensland, Australia), such as possessing a small number of cerarii (≤ eight pairs), oral collar tubular ducts of two sizes, and lacking cerarii on the head; however, it differs from C. melaleucae in having relatively long dorsal setae, each approximately 17–70 µm long, whereas C. melaleucae has relatively short dorsal setae, each approximately 8–16 µm long., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on pages 557-560, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Kanda, S. (1933) Two new species of the genus Pseudococcus from Yokohama and the island of Palau. Annotationes Zoologicae Japonenses, 14, 133 - 138.","Shiraiwa, H. (1935) Studies on mealybugs infesting pear in Japan. Kontyu, 9, 63 - 75.","Shinji, O. (1936) Two new Japanese species of Pseudococcus (Hemiptera-Homoptera). Kontyu, 10, 49 - 51.","Kanda, S. (1941) [On the scientific names of some scale insects]. Kontyu, 15, 24 - 26. [in Japanese]","Paik, W. H. (1978) Illustrated flora and fauna of Korea. Insecta (VI). No. 22. Ministry of Education (Samhwa Publ. Co. Ltd), Soul, 481 pp.","Kawai, S. (2003) Kaigaramushi [Coccoidea]. In: Umeya, K. & Okada, T. (Eds.), Agricultural insect Pests in Japan. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 1 - 1203. [in Japanese]","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Ezzat, Y. M. & McConnell, H. S. (1956) A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A-e 84, 1 - 108.","Williams, D. J. (2004) Mealybugs of southern Asia. The Natural History Museum, London and Southdene SDN. BHD., Kuala Lumpur, 896 pp.","Williams, D. J. (1985) Australian mealybugs. British Museum (Natural History), London, 431 pp."]}

Genus Crisicoccus Ferris 1950 Crisicoccus Ferris 1950: 45. Type species: Dactylopius pini Kuwana 1902. Genus diagnosis (adapted and modified from Williams 2004). Body of adult female normally broadly oval; anal lobe developed, each lobe with an anal lobe bar and bar seta present on ventral surface. Antennae each usually with 7 or 8 segments. Legs well developed, translucent pores usually present (rarely absent) on hind coxae, present also on hind tibiae and sometimes on hind femora. Tarsal digitules knobbed. Circulus normally present. Anterior and posterior ostioles present. Anal ring normal, usually bearing 6 setae. Cerarii numbering 0–17 pairs, each containing 2 conical setae or sometimes reduced to a single conical seta; often cerarii on head and thorax indistinct; preocular cerarii (C 2) absent. Auxiliary setae rarely present except on anal lobe cerarii. Body setae often flagellate, sometimes conical or spiniform. Trilocular pores normal. Multilocular disc pores present, usually confined to venter. Oral rim tubular ducts absent. Oral collar tubular ducts present on venter and sometimes also on dorsum. Remarks. According to Williams (2004), Crisicoccus is considered a convenient genus to include any species possessing well-developed anal lobe bars that do not fit the features of other mealybug genera with anal lobe bars. Crisicoccus is quite similar to Formicococcus Takahashi 1928 and Planococcus Ferris 1950 and so it may be difficult to distinguish these genera. Further morphological and molecular phylogenetic studies of these three genera are greatly needed., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on page 556, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Ferris, G. F. (1950) Atlas of the Scale Insects of North America. Series 5. Vol. 5. The Pseudococcidae (Part I). Stanford University Press, Palo Alto, California, 278 pp.","Kuwana, S. I. (1902) Coccidae (scale insects) of Japan. Proceedings of the California Academy of Sciences, 3, 43 - 98.","Williams, D. J. (2004) Mealybugs of southern Asia. The Natural History Museum, London and Southdene SDN. BHD., Kuala Lumpur, 896 pp.","Takahashi, R. (1928) Coccidae of Formosa (2). Transactions of the Natural History Society of Formosa. Taihoku, 18, 253 - 261."]}

Crisicoccus azaleae (Tinsley 1898) [Japanese common name: Azarea-kona-kaigaramushi] Dactylopius azaleae Tinsley 1898: 319. Pseudococcus azaleae (Tinsley 1898); Fernald 1903: 88 (change of combination). Pseudococcus taxi Kanda 1943: 51. (synonymy by Kawai 1980: 109). Crisicoccus azaleae (Tinsley 1898); Ferris 1953: 305 (change of combination); McKenzie 1967: 125; Kawai 1980: 109; Kawai 2003: 314; Danzig & Gavrilov-Zimin 2015: 200. Planococcus azaleae (Tinsley 1898); Ezzat & McConnell 1956: 63 (change of combination). Crisicoccus taxi (Kanda 1943); Kawai 1972: 6 (change of combination). Material examined. Japan: Tokyo, Akikawa-shi, Nobe, on Rhododendron amagianum, 5.viii.1972, coll. S. Kawai, 3 adult females mounted singly and 2 adult females mounted together on a slide (KTUA); Ibaraki Prefecture, Tsukuba, Kannon-dai, on Rhododendron x pulchrum, 21.iv.2021, coll. J. Tabata, 4 adult females mounted singly (2 ELKU, 2 EUMJ). Updated description Appearance in life. Adult female 3‒4 mm long, dark purple to purple-brown, covered with a white powdery wax. Projections of wax secretion from body margin short and indistinct on cephalothorax, slightly longer on a few segments of posterior part of body. Body contents of this species turn blue-black to dark green in 10% potassium hydroxide solution (Kawai 1980, translated by HT). Slide-mounted adult female (Fig. 4) (n = 9). Body elongate oval, 1.9–3.1 mm long and 0.9–1.9 mm wide; derm membranous; segmentation recognizable but not well developed. Anal lobes well developed, ventral surface with an anal lobe bar and a long apical seta, 154–239 µm long. Antenna 334–432 µm long, with 8 segments and many flagellate setae; subapical segment with 1 fleshy seta, apical segment with 3 fleshy setae. Eyespot present on margin, not associated with discoidal pores. Legs well developed, with many flagellate setae; hind leg measurements (in µm): trochanter + femur 248–320 long; hind tibia + tarsus 283–350; claw 26–36, without a denticle. Ratio of lengths of hind tibia + tarsus to trochanter + femur about 1: 1.04–1.16; ratio of lengths of hind tibia to tarsus 1: 1.79–2.40. Paired setose tarsal digitules present, subequal in length to the minutely knobbed claw digitules. Hind coxa with translucent pores on both surfaces; hind tibia sometimes with translucent pores on posterior surface; hind trochanter, femur and tarsus without translucent pores. Labium 120–152 µm long, shorter than clypeolabral shield. Circulus usually oval but rarely quadrate, located between abdominal segments III and IV or rarely on abdominal segment IV, usually divided by an intersegmental line, 40–113 µm long and 84–120 µm wide. Ostioles present, each with inner edges of lips weakly sclerotized; anterior ostioles each with a total for both lips of 17–29 trilocular pores and 2–7 setae; each posterior ostiole with a total for both lips of 16–44 trilocular pores and 4–7 setae. Anal ring 84–112 µm wide, bearing 6 setae, each seta 104–160 µm long. Cerarii numbering 6–12 pairs, normally present on posterior abdominal segments, rarely present also on thoracic segments. Anal lobe cerarii (C 18) each containing mostly 2 (rarely 3) conical cerarian setae, each 14–22 µm long and about 5–7 µm wide at base; 3–6 auxiliary setae, and a concentration of trilocular pores. Penultimate cerarii (C 17) each containing 2 conical setae, 0 auxiliary setae and a few trilocular pores. Cerarii situated further forward generally each with 2 conical cerarian setae and a few trilocular pores. Dorsum. Setae spiniform, mostly straight, each 7–30 µm long, usually distributed segmentally; longest setae present on head or posterior abdominal segments. Trilocular pores each 3–4 µm wide, evenly distributed. Oral rim tubular ducts and oral collar tubular ducts absent. Discoidal pores each about 2 µm wide, sparsely distributed. Venter. Setae relatively long and flagellate, each 11–140 µm long; setae on head longest. Multilocular disc pores, each 6–9 µm wide, present in medial areas of abdominal segments IV‒IX, arranged in single rows on each posterior area of abdominal segments IV‒ V, 2 or 3 rows on each posterior area of abdominal segments VI –VII, and in a single row on each anterior area of abdominal segments VI –VII but arranged randomly on abdominal segments VIII and IX. Trilocular pores, each of same size as those on dorsum, evenly distributed. Oral collar tubular ducts all of 1 size, each about 2–4 µm wide, present on medial areas of abdominal segments IV–VII and usually forming transverse bands across segments; also relatively slightly stouter tubular ducts present on submarginal to marginal areas of abdominal segments I–VIII and thoracic segments. Discoidal pores, same width as those on dorsum, sparsely present. Host plants in Japan. Ebenaceae: Diospyros kaki (Kawai 1980; 2003); Ericaceae: Rhododendron amagianum (Kawai 1980), Rhododendron macrosepalum (Kawai 1980), Rhododendron spp. (Kawai 2003), Rhododendron x pulchrum; Fabaceae: Albizia julibrissin (Kawai 1980); Fagaceae: Castanopsis cuspidata (Kawai 1980); Magnoliaceae: Magnolia kobus (Kawai 1980); Oleaceae: Ligustrum lucidum (Kawai 1980; 2003); Rosaceae: Eriobotrya japonica (Kawai 1980; 2003), Pyracantha angustifolia (Kawai 1980), Pyrus pyrifolia var. culta (Kawai 1980; 2003); Sapindaceae: Acer buergerianum (Kawai 1980; 2003); Schisandraceae: Illicium anisatum (Kawai 1980); Salicaceae: Salix babylonica var. babylonica (Kawai 1980); Taxaceae: Cephalotaxus harringtonia (Kanda 1943), Taxus cuspidata (Kawai 1980). Remarks. Crisicoccus azaleae is similar to C. pini (Kuwana 1902) in having a small number of cerarii (C. pini in the following morphological characteristics (contrasting characteristics of C. pini are given in parentheses): (i) a circulus usually present between abdominal segments III and IV on the venter (circulus always lacking); (ii) presence of spiniform and mostly straight dorsal setae (dorsal setae are mostly flagellate and slightly curved); and (iii) thoracic segments with a few ventral oral collar tubular ducts (thoracic segments without ventral oral collar tubular ducts). In particular, C. azaleae is characterized by straight, spiniform dorsal setae, which are significantly different from the curved flagellate setae in C. pini. Danzig & Gavrilov-Zimin (2015) stated that “ Crisicoccus azaleae (Tinsley 1898) is very similar, probably conspecific with C. pini ”. However, both species are clearly different and are treated here as wellseparated species., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on pages 562-564, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Tinsley, J. D. (1898) Notes on Coccidae, with descriptions of new species. Canadian Entomologist, 30, 317 - 320. https: // doi. org / 10.4039 / Ent 30317 - 12","Fernald, M. E. (1903) A catalogue of the Coccidae of the world. Bulletin of the Hatch Experiment Station of the Massachusetts Agricultural College, 88, 1 - 360. https: // doi. org / 10.5962 / bhl. title. 8533","Kanda, S. (1943) Two new Pseudococcus (Coccidae) Homoptera from Nippon. Annotationes Zoologicae Japonenses, 22, 49 - 53.","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Ferris, G. F. (1953) Atlas of the Scale Insects of North America. Vol. 6. The Pseudococcidae (Part II). Stanford University Press, Palo Alto, California, 506 pp.","McKenzie, H. L. (1967) Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley, 526 pp. https: // doi. org / 10.1525 / 9780520338227","Kawai, S. (2003) Kaigaramushi [Coccoidea]. In: Umeya, K. & Okada, T. (Eds.), Agricultural insect Pests in Japan. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 1 - 1203. [in Japanese]","Danzig, E. M. & Gavrilov-Zimin, I. A. (2015) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 2: Subfamily Pseudococcinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 619 pp.","Ezzat, Y. M. & McConnell, H. S. (1956) A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A-e 84, 1 - 108.","Kawai, S. (1972) Diagnostic notes and biology of the coccid species occurring on cultivated or wild trees and shrubs in Japan (Homoptera: Coccoidea). Bulletin of the Tokyo-To Agricultural Experiment Station, 6, 1 - 54. [in Japanese]","Kuwana, S. I. (1902) Coccidae (scale insects) of Japan. Proceedings of the California Academy of Sciences, 3, 43 - 98."]}

Spilococcus pacificus (Borchsenius 1949) Pseudococcus pacificus Borchsenius 1949: 119. Paracoccus betulae Borchsenius & Kozarzhevskaya 1966 (synonymy by Danzig 1980: 158). Spilococcus pacificus (Borchsenius 1949); Danzig 1980: 158 (change of combination); Ben-Dov 1994: 491; Suh 2020: 2. Atrococcus pacificus (Borchsenius 1949); Tang 1992: 228 (change of combination); Danzig & Gavrilov-Zimin 2015: 270. Crisicoccus matsumotoi (Shiraiwa 1935); Paik 1978: 186; Paik 2000: 64; Kwon et al. 2003: 403; Suh 2020: 2 (misidentifications). Material examined. South Korea: Busan, on Acer palmatum, 20.v.2001, no collector indicated, 1 adult female mounted singly (APQA); Gunwi (GB), on Pyrus ussuriensis, 13.ix.2012, coll. Suh, S.J., 2 adult females mounted on a slide (APQA). Updated description Appearance in life. Body orange in life (Danzig & Gavrilov-Zimin 2015). Slide-mounted adult female (Fig. 7) (n = 3): Body elongate oval, 2.7–3.3 mm long and 1.4–1.6 mm wide; derm membranous; segmentation recognizable, but not well developed. Anal lobes well developed; dorsal surface of each lobe with a slightly sclerotized area; ventral surface with well-developed anal lobe bar and long apical seta, 208–271 µm long. Antenna 360–450 µm long, with 8 segments and many flagellate setae; subapical segment with 1 fleshy seta and apical segment with 3 fleshy setae. Eyespots present on margin, not associated with discoidal pores. Legs well developed, with many flagellate setae; hind leg measurements (in µm): trochanter + femur 260–338 long; hind tibia + tarsus 280–395; claw 34–40, without a denticle. Ratio of lengths of hind tibia + tarsus to trochanter + femur about 1: 1.01–1.17; ratio of lengths of hind tibia to tarsus 1: 1.55–2.17. Paired setose tarsal digitules present, subequal in length to the minutely knobbed claw digitules. Hind coxa with translucent pores on both surfaces; hind tibia with or without translucent pores on posterior surface; hind trochanter, femur and tarsus without translucent pores. Labium 165–186 µm long, usually same length as clypeolabral shield. Circulus usually quadrate, located between abdominal segment III and IV, 68–170 µm long and 87–125 µm wide. Ostioles present, each with inner edges of lips weakly sclerotized; anterior ostioles each with a total for both lips of 16–29 trilocular pores and 3–5 setae; each posterior ostiole with a total for both lips of 19–49 trilocular pores and 2–5 setae. Anal ring 86–98 µm wide, bearing 6 setae, each seta 110–140 µm long. Cerarii numbering 6–8 pairs, all present on abdominal segments. Anal lobe cerarii (C 18) mostly each containing 2 conical cerarian setae, each seta 12–21 µm long and about 4–7 µm wide at base, 4–7 auxiliary setae and a concentration of trilocular pores. Penultimate cerarii (C 17) each containing 2 conical setae, no auxiliary setae and some trilocular pores. Cerarii situated further forward generally each with 2 conical setae and usually a few trilocular pores. Dorsum. Setae flagellate, each 8–45 µm long, distributed segmentally; longest setae present on head. Trilocular pores each 3–4 µm wide, evenly distributed. Oral collar tubular ducts absent. A few oral rim tubular ducts present on submarginal to medial areas of head, thoracic and abdominal segments. Discoidal pores each about 2–3 µm wide, sparsely distributed. Venter. Setae relatively long and flagellate, each 18–136 µm long; setae on head longest. Multilocular disc pores, each 6–9 µm wide, present in medial areas of abdominal segments IV‒IX; a few also present on head and thoracic segments, arranged in 1 to 3 rows on posterior areas of abdominal segments VI –VII but arranged randomly on abdominal segments VIII and IX. Trilocular pores, each same size as those on dorsum, evenly distributed. Oral collar tubular ducts of 2 sizes present: (i) large-type ducts, each about 3–4 µm in diameter, mostly wider than a trilocular pore, present on marginal to submarginal areas of all abdominal segments and anterior of thorax; and (ii) small-type ducts, each about 1–2 μm in diameter, present on medial areas of abdominal segments IV–VII, forming transverse bands across most posterior abdominal segments. Discoidal pores, each about 1–2 µm wide, sparsely distributed. Host plants in South Korea. Moraceae: Broussonetia kazinoki (Paik 1978; Suh 2020), Rosaceae: Malus pumila (Kwon et al. 2003; Suh 2020), Pyrus pyrifolia var. cultiva (Kwon & Han 2003; Suh 2020), P. ussuriensis (Rosaceae) (Kwon et al. 2003; Suh 2020); and Sapindaceae: Acer palmatum (Paik 1978; Suh 2020). Remarks. In South Korea, this species was first reported as “ C. matsumotoi ” by Paik (1978) and later by Paik (2000), Kwon et al. (2003), and Suh (2020). However, the morphology of the Korean specimens examined in this study, which were identified by South Korean mealybug experts Suh, S.J. and Park, M.J. as “ C. matsumotoi ” is quite different from that of the species currently regarded as C. matsumotoi (= Crisicoccus seruratus) in Japan, the native country of the type specimen. The Korean material could be identified as a member of the genus Spilococcus, namely S. pacificus. The true C. matsumotoi (= C. seruratus) is a well-known significant pest of fruit crops in Japan (Kawai 1980; 2003) and, based on our investigation, is probably absent from South Korea. Thus, this rectification of the misidentifications will be beneficial to agriculture and international plant quarantine. Where mentioned above, Suh (2020) just cited an earlier paper by South Korean researchers; she was not responsible for the misidentifications. In this study, we could not check all the specimens of C. matsumotoi that were used by the South Korean researchers; possibly a few of them could be the true C. matsumotoi (= C. serruratus) from South Korea. However, the morphological characteristics of the Korean specimens we examined in this study are consistent with those of “ C. matsumotoi ” in the key provided by Kwon et al. (2003), such as having some oral rim tubular ducts on the dorsum. We now consider that most or all specimens reported as “ C. matsumotoi ” from South Korea are consistent with S. pacificus. We also believe that the specimens used by other researchers should be re-examined in the future, especially as Suh (2020) cited Kwon & Han (2003) as having recorded S. pacificus on Acer palmatum in Korea, whereas Kwon et al. (2003) mentioned “ C. matsumotoi ” on the same host plant. All the Korean specimens examined in this study have oral-rim tubular ducts on the dorsum and anal lobe bars on the ventral surfaces of the anal lobes. According to Kaydan & Szita (2017), the species should be transferred either to the genus Maconellicoccus Ezzat or to Paracoccus Ezzat & McConnell (probably, to the genus Paracoccus with minor generic diagnostic modifications); however, in the present study, none of the type specimens were examined so the species is left in Spilococcus for now., Published as part of Tanaka, Hirotaka & Kamitani, Satoshi, 2022, Review ofthegenus CrisicoccusFerris (Hemiptera: Coccomorpha: Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935), pp. 555-572 in Zootaxa 5209 (5) on pages 568-570, DOI: 10.11646/zootaxa.5209.5.3, http://zenodo.org/record/7340959, {"references":["Borchsenius, N. S. (1949) [Insects Homoptera. suborders mealybugs and scales (Coccoidea). Family mealybugs (Pseudococcidae). Vol. VII.]. Fauna SSSR. Zoologicheskii Institut Akademii Nauk SSSR, New Series, 38, 1 - 382. [in Russian]","Danzig, E. M. (1980) [Coccoids of the Far East USSR (Homoptera, Coccinea) with phylogenetic analysis of scale insects fauna of the world]. Nauka, Leningrad, 367 pp. [in Russian]","Ben-Dov, Y. (1994) A systematic catalogue of the mealybugs of the world (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with data on geographical distribution, host plants, biology and economic importance. Intercept Limited, Andover, 686 pp.","Suh, S. J. (2020) Host plant list of the scale insects (Hemiptera: Coccomorpha) in South Korea. Insecta Mundi, 0757, 1 - 26.","Tang, F. T. (1992) [The Pseudococcidae of China.] Chinese Agricultural Science Technology Press Beijing, 768 pp. [in Chinese with English abstract]","Danzig, E. M. & Gavrilov-Zimin, I. A. (2015) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 2: Subfamily Pseudococcinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 619 pp.","Shiraiwa, H. (1935) Studies on mealybugs infesting pear in Japan. Kontyu, 9, 63 - 75.","Paik, W. H. (1978) Illustrated flora and fauna of Korea. Insecta (VI). No. 22. Ministry of Education (Samhwa Publ. Co. Ltd), Soul, 481 pp.","Paik, J. C. (2000) Economic Insects of Korea 6, Homoptera (Coccinea), Insecta Koreana Supplement 13. National Institute of Agricultural Science and Technology, Seoul, 193 pp.","Kwon, G. M. & Han, M. J. (2003) [Scale insects (Stenorrhyncha) occurred on fruit trees in Korea]. Korean Journal of Applied Entomology, 42 (4), 279 - 288. [in Korean]","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Kawai, S. (2003) Kaigaramushi [Coccoidea]. In: Umeya, K. & Okada, T. (Eds.), Agricultural insect Pests in Japan. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 1 - 1203. [in Japanese]","Kaydan, M. B. & Szita, E. (2017) Mealybugs (Hemiptera: Coccomorpha: Pseudococcidae) with oral rim ducts; description of a new genus and species from Turkey, and discussion of their higher classification within the Pseudococcidae. Zootaxa, 4227 (3), 444 - 450. https: // doi. org / 10.11646 / zootaxa. 4227.3.10"]}

Tanaka, Hirotaka, Kamitani, Satoshi (2022): Review ofthegenus CrisicoccusFerris (Hemiptera:Coccomorpha:Pseudococcidae) in Japan with description of a new species, and the identity of a South Korean mealybug misidentified as Crisicoccus matsumotoi (Shiraiwa 1935). Zootaxa 5209 (5): 555-572, DOI: https://doi.org/10.11646/zootaxa.5209.5.3

Formicococcus kawaii sp. nov. (Fig. 2) [Japanese common name: Oo-take-kona-kaigaramushi] Material examined. Holotype adult female: Japan, / Tokyo, Minato-ku, / Minami-Aoyama, / Aoyama cemetery / on Pleioblastus chino / 7.vi.1972 / coll. S. Kawai, mounted singly (KTUA); Paratype: same data as holotype, 1 adult female mounted singly (KTUA)., Published as part of Tanaka, Hirotaka, 2022, A taxonomic study of three mealybug species (Hemiptera: Coccomorpha: Pseudococcidae) infesting dwarf bamboo in Japan, with description of a new species, pp. 334-346 in Zootaxa 5178 (4) on page 339, DOI: 10.11646/zootaxa.5178.4.2, http://zenodo.org/record/7031678

Paraputo kaiensis (Kanda 1932) (Fig. 3) [Japanese common name: Miyama-kona-kaigaramushi] Pseudococcus kaiensis Kanda 1932: 387. Kaicoccus kaiensis (Kanda 1932); Takahashi 1958: 5; Kawai 1980: 113. Dysmicoccus kaiensis (Kanda 1932); Danzig 1971: 367. Paraputo kaiensis (Kanda 1932); Danzig & Gavrilov-Zimin 2015: 29. Material examined. JAPAN: Tochigi prefecture, Nikko-shi, on Bambusoideae sp. (dwarf bamboo), 21.vii.1934, coll. S. Kanda, 1 adult female mounted singly (OMNH); Gunma prefecture, Tsumagoi-mura, Mt. Yunomaru, on Bambusoideae sp. (dwarf bamboo), 22.vii.1960, coll. S. Kanda, 2 adult females mounted singly (OMNH); Gunma prefecture, Azuma-gun, Kusatsu-cho, Mt. Shirane, on Bambusoideae sp. (dwarf bamboo), 1.viii.1960, coll. S. Kanda, 2 adult females mounted on 1 slide (OMNH); Tochigi prefecture, Mt. Nasu, on Bambusoideae sp. (dwarf bamboo), 13.vi.1976, coll. S. Kawai, 6 adult females mounted singly (KTUA)., Published as part of Tanaka, Hirotaka, 2022, A taxonomic study of three mealybug species (Hemiptera: Coccomorpha: Pseudococcidae) infesting dwarf bamboo in Japan, with description of a new species, pp. 334-346 in Zootaxa 5178 (4) on page 342, DOI: 10.11646/zootaxa.5178.4.2, http://zenodo.org/record/7031678, {"references":["Kanda, S. (1932) A new species of the genus Pseudococcus. Annotationes Zoologicae Japonenses. Tokyo, 13, 387 - 390.","Takahashi, R. (1958) Key to the genera of Pseudococcidae in Japan, with descriptions of three new genera and two new species. Bulletin of the Osaka (Perfecture) University, 7 (1957), 1 - 8.","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Danzig, E. M. (1971) New and little-known species of mealy-bugs (Homoptera, Coccoidea, Pseudococcidae) from the Far East of USSR. Entomologicheskoe Obozrenye, 50, 366 - 391.","Danzig, E. M. & Gavrilov-Zimin, I. A. (2015) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 2: Subfamily Pseudococcinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 619 pp."]}

Formicococcus bambusiphilus (Takahashi 1958) stat. rev. and comb. nov. (Fig. 1) [Japanese common name: Take-kona-kaigaramushi] Ferrisicoccus bambusiphilus Takahashi 1958: 7; Kawai 1972: 7; Kawai 1980: 114. Ferrisicoccus angustus Ezzat & McConnell 1956; Ben-Dov 1999: 380 (misidentification). Dysmicoccus angustus (Ezzat & McConnell 1956); Danzig 1978: 8 (misidentification). Paraputo angustus (Ezzat & McConnell 1956); Danzig & Gavrilov-Zimin 2015: 29 (misidentification). Material examined. Lectotype (here designated): Ferrisicoccus / bambusiphilus / Takah. Type / 19.VIII.1952 / Tokyo, Japan / R. Takahashi. Host: Bamboo. 1 adult female on a slide together with 9 paralectotypes; the lectotype is the last individual on the right in the upper part of the slide (SEHU); Paralectotype: same data as lectotype; examined paralectotype is the specimen on the left below the lectotype (SEHU). Other material studied: JAPAN: Kanagawa prefecture, Atsugi-shi, Nanasawa, on Pleioblastus chino, 25.vii.1975, coll. S. Kawai, 2 adult females mounted singly (KTUA); Tokyo, Chiyoda-ku, Imperial palace, on Pleioblastus chino, 24.vi.1998, coll. S. Kawai, 4 adult females mounted singly (KTUA)., Published as part of Tanaka, Hirotaka, 2022, A taxonomic study of three mealybug species (Hemiptera: Coccomorpha: Pseudococcidae) infesting dwarf bamboo in Japan, with description of a new species, pp. 334-346 in Zootaxa 5178 (4) on page 336, DOI: 10.11646/zootaxa.5178.4.2, http://zenodo.org/record/7031678, {"references":["Takahashi, R. (1958) Key to the genera of Pseudococcidae in Japan, with descriptions of three new genera and two new species. Bulletin of the Osaka (Perfecture) University, 7 (1957), 1 - 8.","Kawai, S. (1972) Diagnostic notes and biology of the coccid species occurring on cultivated or wild trees and shrubs in Japan (Homoptera: Coccoidea). Bulletin of the Tokyo-To Agricultural Experiment Station, 6, 1 - 54. [in Japanese]","Kawai, S. (1980) Scale Insects of Japan in Colors. Zenkoku Noson Kyoiku Kyokai, Tokyo, 455 pp. [in Japanese]","Ezzat, Y. M. & McConnell, H. S. (1956) A classification of the mealybug tribe Planococcini (Pseudococcidae: Homoptera). Bulletin of the Maryland Agriculture Experiment Station, A-e 84, 1 - 108.","Ben-Dov, Y. (1999) A note on Ferrisicoccus Ezzat & McConnell (Hem., Coccoidea, Pseudococcidae). Bulletin de la Societe entomologique de France, 104 (4), 380. https: // doi. org / 10.3406 / bsef. 1999.16599","Danzig, E. M. (1978) Scale insect fauna of South Sakhalin and Kunashir. Trudy Biologo-Pochvennogo, Akademii Nauk SSR, Vladivostok, 50, 3 - 23.","Danzig, E. M. & Gavrilov-Zimin, I. A. (2015) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 2: Subfamily Pseudococcinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 619 pp."]}

Dysmicoccus kunaw Tanaka sp. nov. (Japanese common name: Tanpopo-kona-kaigaramushi) Material examined. Holotype: JAPAN, Hokkaido, / Kamikawa-gun, / Pippu-cho, / Kamikawa Agricultural / Experiment Station, / on Taraxacum officinale, / 9.vii.2020, / coll. D. Sasaki; adult female mounted singly (ELKU). Paratypes: same data as for holotype; 9 adult females mounted singly (4 ELKU, 5 EUMJ). Paratype used for molecular phylogenetic analysis: same data as for holotype; 1 adult female (1 ELKU, Genbank Accession No. ON533755 for COI, ON527952 for 18S, ON527954 for 28S D2). Description (n = 11). Live adult female: Feeding on the roots of the host plant and secreting white powdery wax on all body surfaces (Fig. 2); the mealybugs are attended by the ant Lasius flavus Fabricius (Hymenoptera: Formicidae). Slide-mounted adult female (Fig. 3): Body elongate oval, 3.1 (2.4–3.1) mm long and 2.0 (1.5–2.2) mm wide; derm membranous; segmentation not well-developed. Anal lobes clearly evident but not prominent, dorsal and ventral surfaces of each lobe without sclerotised areas and ventral surface with long apical seta, 160–225 (150– 235) µm long. Antenna 439–444 (375–447) µm long, with 7 or 8 segments (usually 8) and many flagellate setae; subapical segment with 1 (0 or 1) fleshy seta and apical segment with 3 (2–4) fleshy setae. Eyes present on margin, not associated with discoidal pores. Legs well-developed, with many flagellate setae; hind trochanter + femur 333–344 (295–348) µm long; hind tibia + tarsus 330–334 (295–340) µm long; claw 40–41 (36–43) µm long without a denticle. Ratio of lengths of hind tibia + tarsus: trochanter + femur about 1: 1.0 (0.9–1.0); ratio of lengths of hind tibia to tarsus 1: 1.8–2.0 (1.7–2.1). Single or paired setose tarsal digitules present, subequal in length to minutely knobbed claw digitules. Hind legs without translucent pores. Labium 120 (120–136) µm long, shorter than clypeus. Circulus absent. Ostioles present, each with inner edges of lips weakly sclerotised; anterior ostioles each with a total for both lips of 22–28 (12–28) trilocular pores and 5 or 6 (1–6) setae; each posterior ostiole with a total for both lips of 42 or 43 (16–47) trilocular pores and 1 or 2 (0–4) setae. Anal ring 90 (78–101) µm wide, with 2 rows of cells, bearing 7 (6 or 7, usually 6) setae, each seta 110–161 (60–170) µm long. Cerarii numbering 15–17 (14–17) pairs. Anal lobe cerarii mostly each containing 2 slender conical cerarian setae, each seta 16–20 (15–20) µm long and about 6–7 (6–8) µm wide at base, also 10–13 (7–16) auxiliary setae and a concentration of trilocular pores. Penultimate cerarii each containing 2 long, stout flagellate cerarian setae and many auxiliary setae. Cerarii situated further forward generally each with 1–3 stout long, flagellate cerarian setae and auxiliary setae; boundary of each cerarius on anterior thoracic segments and head obscure and sometimes difficult to distinguish. Dorsum. Setae long and flagellate, each 49–132 (40–162) µm long, distributed segmentally. Trilocular pores, each 3–4 µm wide, evenly distributed. Oral collar tubular ducts of 2 sizes: (i) large-type ducts, each about 3–4 µm in diameter, mostly each same width as a trilocular pore, present on all segments of dorsum, forming transverse bands across most segments; and (ii) small-type ducts, each about 1–2 μm in diameter, relatively sparse on all segments, intermixed with large-type ducts. Discoidal pores, each 2–3 µm wide, sparsely distributed. Multilocular disc pores each 7–9 (7–10) µm wide, present on medial area of abdominal segments as follows: segment I, 0 (0); II, 0 (0 or 1); III, 1 (0 or 1); IV, 5 (0–6); V, 0 (0–14); VI, 2 (2–10); VII, 7 (1–11); and VIII, 0 (0 or 1). Venter. Setae long and flagellate, each 64–160 (25–194) µm long, distributed segmentally, longest on medial area of head or posterior abdominal segments. Multilocular disc pores, each 7–9 (7–11) µm wide, present on medial area of all segments of venter between head and abdominal segment IX. Trilocular pores, same width as on dorsum, evenly distributed. Oral collar tubular ducts of 2 sizes: (i) large-type ducts, each about 3–4 µm in diameter, mostly same width as trilocular pores, present on all segments of venter, forming transverse bands across most segments; and (ii) small-type ducts, each about 1–2 μm in diameter, relatively sparse between mesothorax and abdominal segments VII, intermixed with large-type ducts. Discoidal pores, same width as on dorsum, sparsely present. Host plants. Roots of Taraxacum officinale (Asteraceae). Remarks. Dysmicoccus kunaw resembles D. trispinosus (Hall 1923) described from Egypt, and D. furcillosus Williams 2004 described from India, in having a slightly rounded body, long flagellate body setae, and long flagellate cerarian setae. However, D. kunaw differs from them as follows (contrasting character states in D. trispinosus and D. furcillosus in parentheses): (i) each anal lobe cerarius with two conical cerarian setae (anal lobe cerarian setae not conical, but long and stout setose); (ii) abdominal segments of dorsum with a considerable number of multilocular pores, also on head and thoracic segments of venter (multilocular disc pores absent from dorsum, and head and thoracic segments of venter); and (iii) ventral oral collar tubular ducts of two types (only one type of ventral duct present in D. trispinosus, and three types in D. furcillosus). The molecular phylogenetic analysis placed D. kunaw within the clade of the subfamily Pseudococcinae (Fig. 1). The new species formed a clade with Paraputo kaiensis (Kanda 1932), although this was not well supported (UFBoot = 84) and showed relatively long branch lengths. This clade was sister to a clade including species of Dysmicoccus, Erium, Palmicultor and Trionymus. A clade with Pseudococcus cryptus Hempel 1918 and P. jackbeardsleyi Gimpel & Miller 1996 was placed outside the clade containing D. kunaw. The molecular analysis does not support the generic designation of the new species in this study, which is based on adult female morphological characteristics. The analysis also indicated that D. kunaw was not related to the type species of Dysmicoccus, the neotropical species D. brevipes (Cockerell 1893). Dysmicoccus is known to be a polyphyletic group (Choi & Lee 2022). Although the genus Dysmicoccus needs revision, we tentatively place D. kunaw in Dysmicoccus based on the current morphological classification. Relationship with ants. The type series of D. kunaw was tended by the ant Lasius flavus Fabricius 1782 (Hymenoptera: Formicidae) and was found inside ant-made carton shelters. Dysmicoccus kunaw may have a close relationship with this and other ant species belonging to the genus Lasius. Etymology. The specific epithet “ kunaw ” is an Ainu noun that refers to a dandelion, a forked-stem adonis, or a goddess of mist in Ainu mythology who was transformed into a flower by the curse of her husband, Hoinu, the marten god. The Ainu are the local indigenous people of Hokkaido Island. The epithet is used as a noun in apposition.

Genus Dysmicoccus Ferris, 1950 Dysmicoccus Ferris 1950: 53. Type species: Dactylopius brevipes Cockerell 1893. Parkermicus Khalid & Shafee 1988: 31. Type species: Parkermicus polyanosetosus Khalid and Shafee 1988. Diagnosis (adapted from Tanaka & Kamitani 2021). Body of adult female elongate to broadly oval. Anal lobes usually developed, either membranous or sclerotised, each lobe bearing a normal apical seta. Ventral margins of abdominal segments anterior to anal lobes always membranous. Antennae each with 6–8 segments. Legs well developed; translucent pores present or absent; tarsal digitules usually knobbed. Claw without a denticle. Cerarii numbering 6–17 pairs. Auxiliary setae present, at least in anal lobe cerarii. Anal lobe cerarii each bearing either 2 cerarian setae or as many as 8, the setae usually conical; sometimes conical setae replaced by flagellate setae but cerarii always recognisable by concentrations of trilocular pores. Anal ring normally situated at apex of abdomen, usually bearing 6 setae.Anterior and posterior ostioles present. Dorsal setae variously shaped. Ventral setae flagellate. Trilocular pores present on both dorsal and ventral surfaces. Multilocular pores usually present, at least on venter. Quinquelocular pores always absent. Oral collar tubular ducts usually present. Oral rim tubular ducts always absent. Discoidal pores present, sometimes large, occasionally present next to each eye. Remarks. The molecular phylogenetic analysis on mealybugs conducted in this study showed that the genus Dysmicoccus is not a simple monophyletic group but forms a single large clade with several other genera, such as Pseudococcus Westwood 1840, Trionymus Berg 1899, Paraputo Laing 1929, etc. (Fig. 1), so the current definition of Dysmicoccus is probably arbitrary. Further molecular and morphological studies on this genus are greatly needed. Key to adult females of Dysmicoccus species in Japan 1(0) Cerarii numbering fewer than 13 pairs.................................................................... 2 - Cerarii numbering more than 14 pairs (usually 17 pairs)....................................................... 3 2(1) Anal lobe cerarius containing more than 10 auxiliary setae. Dorsum with multilocular pores. Body narrow and parallel sided, with ratio of maximum body length: width 1: 2.8–3.5.......................... D. bunagaya Tanaka & Kamitani 2021 - Anal lobe cerarius containing fewer than 10 auxiliary setae. Dorsum without multilocular pores. Body relatively broad and not parallel-sided, with ratio of maximum body length: width 1: 1.7–2.0....................... D. boninsis (Kuwana 1909) 3(1) Eyes not associated with discoidal pores................................................................... 4 - Eyes associated with discoidal pores...................................................................... 5 4(3) Dorsum with multilocular pores. Hind femora and tibiae without translucent pores. Venter of each anal lobe without narrow irregular sclerotised bar................................................................................ 6 - Dorsum without multilocular pores. Hind femora and tibiae with translucent pores. Venter of each anal lobe with narrow irregular sclerotised bar............................................................ D. wistariae (Green 1923) 5(3) Abdominal segments VII and VIII with dorsal setae longer than those elsewhere, each almost as long as an anal ring seta................................................................................. D. brevipes (Cockerell 1893) - Abdominal segments VII and VIII with dorsal setae short, each much shorter than an anal ring seta............................................................................................... D. neobrevipes (Beardsley 1959) 6(4) Cerarian setae situated on penultimate cerarii and further forward cerarii stout, long and flagellate. D. kunaw Tanaka sp. nov. - Cerarian setae situated on penultimate cerarii and further forward cerarii short and conical..... D. walkeri (Newstead 1891), Published as part of Tanaka, Hirotaka, Sasaki, Daisuke, Choi, Jinyeong, Husnik, Filip & Kamitani, Satoshi, 2022, Two new species of mealybugs (Hemiptera: Coccomorpha: Pseudococcidae) from Japan, pp. 306-318 in Zootaxa 5168 (3) on pages 307-309, DOI: 10.11646/zootaxa.5168.3.3, http://zenodo.org/record/6882716, {"references":["Khalid, M. & Shafee, S. A. (1988) A new genus of Pseudococcidae (Homoptera: Coccoidea) from Manipur, India. Indian Journal of Systematic Entomology, 5, 31 - 33.","Choi, J. & Lee, S. (2022) Higher classification of mealybugs (Hemiptera: Coccomorpha) inferred from molecular phylogeny and their endosymbionts. Systematic Entomology, 47 (2), 354 - 370. https: // doi. org / 10.1111 / syen. 12534","Tanaka, H. & Kamitani, S. (2021) Two new species of Coccomorpha (Hemiptera: Sternorrhyncha) collected from Japanese silver grass, Miscanthus sinensis (Poaceae) in Okinawa Island, Japan. Zootaxa, 4941 (4), 569 - 579. https: // doi. org / 10.11646 / zootaxa. 4941.4.6","Kuwana, S. I. (1909) Coccidae of Japan (IV). A list of Coccidae from the Bonin Islands (Ogasawarajima), Japan. Journal of the New York Entomological Society, 17, 158 - 164.","Green, E. E. (1923) Observations on British Coccidae - VIII. Entomologist's Monthly Magazine, 59, 211 - 218.","Beardsley, J. W. (1959) On taxonomy of pineapple mealybugs in Hawaii, with a description of a previously unnamed species (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, 17, 29 - 37.","Newstead, R. (1891) On some new or little known Coccidae found in England. Entomologist's Monthly Magazine, 26, 164 - 166."]}

Phenacoccus miruku Tanaka & Choi sp. nov. (Japanese common name: Miroku-kona-kaigaramushi) Material examined. Holotype: JAPAN, / Okinawa prefecture, / Okinawa Is. Ogimi-son, / on Bidens pilosa / var. radiata, / 26.vi.2021, / coll. J. Choi; adult female mounted singly (ELKU) . Paratypes: same data as for holotype; 13 adult females mounted singly (6 ELKU, 7 EUMJ). Paratype used for molecular phylogenetic analysis: same data as for holotype; 1 adult female (1 ELKU, Genbank Accession No. ON533756 for COI, ON527953 for 18S, ON527955 for 28S D2). Description (n = 15). Live adult female: Body elongate oval, yellowish, with a thin layer of white powdery wax, very short lateral filaments around entire body margin, and caudal filaments slightly longer and thicker than lateral filaments. At the oviposition stage, the adult female produces a rather loose ovisac of fine wax filaments and lays eggs in it (Fig. 4). Slide-mounted adult female (Fig. 5): Body elongate oval, 1.8 (1.5–2.1) mm long and 1.0 (0.8–1.2) mm wide; derm membranous; segmentation not well developed. Anal lobes evident but not prominent, dorsal and ventral surfaces of each lobe without sclerotised areas and ventral surface with long apical seta, 150 (105–180) µm long. Antenna 327 (304–387) µm long, with 8 or 9 segments (usually 9) and many flagellate setae; subapical segment with 1 fleshy seta and apical segment with 2 (0–3) fleshy setae. Eyes present on margin. Legs well-developed, with many flagellate setae; hind trochanter + femur 243–252 (205–269) µm long, hind tibia + tarsus 310–318 (268–331) µm long; claw 24–30 (22–33) µm long, with a denticle. Ratio of lengths of hind tibia + tarsus: trochanter + femur 1: 1.3 (1.2–1.3); ratio of lengths of hind tibia to tarsus 1: 2.4–2.6 (1.8–2.6). Paired setose tarsal digitules present (most of them partially broken in examined specimens), subequal in length to slightly knobbed claw digitules. Hind legs without translucent pores. Labium about 146 (108–146) µm long, shorter than clypeus. Circulus present or absent, if present located in posterior part of abdominal segment III, 24 (4–51) µm long and 33 (6–122) µm wide. Ostioles present, each with inner edges of lips weakly sclerotised; anterior ostioles each with a total of 16 or 17 (6–23) trilocular pores and 3 or 4 (0–5) setae on both lips; each posterior ostiole with a total of 23–29 (6–35) trilocular pores and 2 or 3 (0–5) setae on both lips. Anal ring 72 (63–88) µm wide, bearing 6 (4–6) setae, each seta 100–112 (74–128) µm long. Cerarii numbering 18 (17 or 18) pairs. Anal lobe cerarii mostly each containing 2 slender conical setae, each seta 14 (8–15) µm long and about 3 (2–5) µm wide at base, also 0 or 1 (0–3) auxiliary setae and a concentration of trilocular pores. Penultimate cerarii each containing 2 slender conical setae but without auxiliary setae. Cerarii situated further forward mostly each with 2 (1–4) conical slender setae only, those cerarii containing 4 slender conical setae mostly located on head. Dorsum. Setae conical to lanceolate, each 3–13 (2–15) µm long, distributed evenly. Trilocular pores, each 3–4 µm wide, evenly distributed. Oral collar tubular ducts of 1 size, each about 2–4 µm in diameter, present on marginal areas of each abdominal segment, and rarely present on marginal areas of head and thoracic segments. Discoidal pores, each 1–2 µm wide, sparsely distributed. Multilocular disc pores each 6–9 µm wide, present on marginal areas of abdominal segments as follows: segment I, 0 (0); II, 0 (0); III, 0 (0 or 1); IV, 0 (0–2); V, 0 (0–3); VI, 0 (0–3); VII, 0 (0–2); and VIII, 0 (0 or 1). Venter. Setae of 2 types: (i) conical to lanceolate setae, each about 2–10 µm long, present on marginal areas of head, thoracic segments and anterior abdominal segments; and (ii) relatively long and slender flagellate setae, each 15–88 (10–109) µm long, present on medial areas of body and marginal areas of posterior abdominal segments, longest on medial area of head. Multilocular disc pores, each 7–9 µm wide, mostly present in medial area of abdominal segments III–IX, occasionally a few also present on marginal areas of abdominal segments. Trilocular pores, same size as on dorsum, evenly distributed. Quinquelocular pores, each 6–7 (3–7) µm wide, present on medial area between lateral mouthparts and abdominal segment III, but occasionally restricted to an area lateral to mouthparts. Oral collar tubular ducts of 1 type, each about 2–4 µm in diameter, present on medial area between prothoracic segment and abdominal segment IX, and on marginal to submarginal areas of posterior abdominal segments, forming transverse bands across all abdominal segments; ducts in marginal areas usually larger; ducts gradually changing in size so cannot be divided into two distinct size types. Discoidal pores, same width as those on dorsum, sparsely present. Host plants. On stems and roots of Bidens pilosa var. radiata (Asteraceae). Remarks. Phenacoccus miruku resembles P. sisymbriifolium Granara de Willink in Granara de Willink & Szumik 2007, described from Uruguay, in having occasional multilocular pores on the marginal areas of the dorsal abdomen, greatly varied conical-to-lanceolate dorsal setae, and conical-to-lanceolate ventral setae on the marginal areas of the venter. However, P. miruku differs from P. sisymbriifolium as follows (contrasting character states in P. sisymbriifolium in parentheses): (i) quinquelocular pores absent in area anterior to mouthparts (ca. 32 quinquelocular pores present in area anterior to mouthparts); (ii) quinquelocular pores absent from abdominal segments IV‒VII (pores present on abdominal segments IV–VII); (iii) translucent pores absent from hind tibiae (some pores present on hind tibiae); and (iv) circulus present or absent on venter, if present, oval (circulus always present, anvil-shaped). Phenacoccus miruku also resembles P. similis Granara de Willink 1983, described from Argentina. However, it differs from the latter as follows (contrasting character states in P. sisymbriifolium in parentheses): (i) quinquelocular pores absent in area anterior to mouthparts (some pores present in area anterior to mouthparts); (ii) translucent pores absent from hind tibiae (some pores present on hind tibiae); and (iii) circulus present on or absent from venter, if present, oval (circulus always present, anvil-shaped). In the phylogenetic tree (Fig. 1), P. miruku was placed within the clade of the subfamily Phenacoccinae. The new species was nested within a clade including both Neotropical and Nearctic species, such as P. manihoti MatileFerrero 1977, P. parvus Morrison 1924, P. peruvianus Granara de Willink in Granara de Willink and Szumik 2007, P. solani Ferris 1918 and P. solenopsis Tinsley 1898. Phenacoccus miruku was sister to a clade containing P. manihoti and P. peruvianus, although this was not well supported (UFBoot = 91). Phenacoccus parvus was placed outside the clade including P. miruku. The molecular analysis does not support the generic designation of this new species in this study, which is based on adult female morphological characteristics. The analysis also indicated that P. miruku is not related to the type species of Phenacoccus, P. aceris (Signoret, 1875). However, Phenacoccus is known to be a polyphyletic group (Choi & Lee 2022). Although this genus needs revision, we tentatively place the new species in Phenacoccus based on the current morphological classification. Based on morphological and molecular evidence, P. miruku is related to Neotropical and Nearctic species, P. manihoti, P. parvus, P. peruvianus, P. sisymbriifolium, P. similis, P. solani and P. solenopsis, so it is possible that P. miruku might have been introduced from the Neotropical or Nearctic regions. It is necessary, therefore, to pay close attention to the current distribution and expansion trend of this species on Okinawa Island and in other regions of Japan. Etymology. The specific epithet “ miruku ” is an Okinawan noun that refers to a visiting god in the mythology of the Japanese Southwest Islands that is sometimes thought to be the same being ethnologically as Miroku Bosatsu (Maitreya Bodhisattva) in Buddhism. The epithet is used as a noun in apposition.