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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., 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 313-316, DOI: 10.11646/zootaxa.5168.3.3, http://zenodo.org/record/6882716, {"references":["Granara de Willink, M. C. & Szumik, C. (2007) Phenacoccinae de Centro y Sudamerica (Hemiptera: Coccoidea: Pseudococcidae): Sistematica y Filogenia. Revista de la Sociedad Entomologica Argentina, 66 (1 - 2), 29 - 129.","Granara de Willink, M. C. (1983) Tres especies de Phenacoccus de la Provincia de Tucuman, Argentina (Hom. Pseudococcidae). Acta Zoologica Lilloana, 37, 59 - 64.","Morrison, H. (1924) The Coccidae of the Williams Galapagos Expedition. Zoologica, 5, 143 - 152. https: // doi. org / 10.5962 / p. 190338","Ferris, G. F. (1918) The California species of mealy bugs. University Series. Stanford University Publications, Palo Alto, California, 78 pp. https: // doi. org / 10.5962 / bhl. title. 22913","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","Signoret, V. (1875) Essai sur les cochenilles ou gallinsectes (Homopteres - Coccides), 15 e partie. Annales de la Societe entomologique de France, Serie 5, 5, 305 - 352.","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"]}

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."]}

List of Euplotes strains and populations analyzed in this paper, and all their bacterial symbionts. Additional fluorescent in situ hybridizations. Formal descriptions of new bacterial taxa. Synopsis of the known bacterial symbionts of Euplotes.