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Genus Compressalges Dubinin, 1950 Type species: Compressalges nipponiae Dubinin, 1950, by original designation. This genus includes only the type species. Within the family Caudiferidae, this genus clearly differs from two other genera, Caudifera Gaud & Mouchet, 1959 and Semicaudifera Gaud & Atyeo, 1996, in having the following features. In males of Compressalges, the opisthosoma is strongly narrowed posteriorly and with a pair of elongated opisthosomal lobes, the ventral side of opisthosoma has long epimerites IVa and adanal apodemes, and legs IV are hypertrophied; in females, the external copulatory tube is absent and the copulatory opening is situated terminally between setae ps1 (Dubinin 1950; Gaud & Atyeo 1996)., Published as part of Waki, Tsukasa & Shimano, Satoshi, 2022, Redescription of two parasitic feather mites sampled from the last two Crested Ibises, Nipponia nippon (Temminck, 1835) (Pelecaniformes: Threskiornithidae) lived in Japan, pp. 136-150 in Zootaxa 5116 (1) on page 138, DOI: 10.11646/zootaxa.5116.1.7, http://zenodo.org/record/6364386, {"references":["Dubinin, V. B. (1950) Features of the structures of the fastening apparatus of the feather mite Compressalges nipponiae V. Dubinin, gen. and sp. nov. Proceedings of the USSR Academy of Sciences, 70, 537 - 540. [in Russian]","Gaud, J. & Mouchet, J. (1959) Acariens plumicoles (Analgesoidea) parasites des oiseaux du Cameroun III. Dermoglyphidae. Annales de Parasitologie humaine et comparee, 34, 149 - 208. https: // doi. org / 10.1051 / parasite / 1959341149","Gaud, J. & Atyeo, W. T. (1996) Feather mites of the world (Acarina: Astigmata): The supraspecific taxa. Musee royal de l'Afrique centrale, Annales, Sciences zoologiques, 277 (Pt. 1 & 2), 1 - 193 (text) & 1 - 436 (illustrations)."]}

Freyanopterolichus nipponiae Dubinin, 1953 [Japanese name: Toki-enban-umoudani] (Figs. 7–10) Freyanopterolichus nipponiae Dubinin, 1953: 292–294, fig. 124; Gaud & Atyeo 1996: 93, fig. 314; Waki & Shimano 2020: 1–8, fig.1; Kuroki et al. 2020, fig.2. Material examined. 4 females and 4 males (MPM Coll. No. 21821) from feathers of Nipponia nippon (Temminck, 1835) (Threskiornithidae), Japan, Niigata Prefecture Sado Island, Ministry of the Environment, Sado Japanese Crested Ibis Conservation Center, 21, August 1993 –4, October 1994, feather coll. Dr. Yoshinori Kaneko. Description. FEMALE (Figs. 7A–B, 9A–D, 10A–B, E, G). Dorsal idiosoma. Subcapitulum trapezoidal, length 77–120, width at base 91–139. Body length including gnathosoma 438–490. Idiosoma transversally ovate, slightly wider than long, strongly flattened dorsoventrally, length 386–445 and greatest width 386–445, width including lateral membranes 467–509. Prodorsal shield large, occupying entire surface of prodorsum, roughly triangular, with two pairs of angular lateral incisions, length 79–97, greatest width 228–249, (Fig. 7A). Setae vi spiculiform, situated near anterior end of prodorsal shield. Setae si and se situated at posterior margin of this shield, setae si narrowly lanceolate, and setae se long sword-shaped, separated by 104–108. Scapular shields represented by small curved sclerites. Lateral margins of hysterosoma with narrow membranes stretching from sejugal furrow to bases of setae f2; length of hysterosoma 310–330. Hysteronotal shield covering most part of hysterosoma, wider than long, anterior margin straight, length 278–290, greatest width 318–457, most surface with polygonal net-work pattern (Fig. 10A), ovate median area near posterior margin with small pit-like lacunae (Fig. 10B). Seta c2 narrowly lanceolate, situated on soft tegument near lateral margins of propodosoma. Seta cp long whip-like, situated marginally slightly posterior to bases of setae c2. Setae d2, e2, f2, h2, h3, ps1, ps2 situated near lateral margins of hysterosoma. Setae d2 filiform. Setae e2, f2, ps1 dilated, narrowly lanceolate, macrosetae h2 and h3 dilated along all their length, gradually attenuate to apices. Seta ps2 short filiform. Hysteronotal gland opening gl posteromesal from setae e2. Cupules ip close tosetae f2, cupules im indistinct. Distance between setae and hysteronotal gland openings: si:si 53–64, se:se 104–118, c1:d1 78–89, d1:gl 65–95, d2:gl 93–120, e1:gl 56–68, e1:h1 89–101, h1:h1 114–154, h2:h2 219–247, h3:h3 148–160, ps1:ps1 103–122. Length of dosal setae: vi 11–15, si 78–113, se 156–158, c1 12–23, c2 70–85, cp 126–156, d1 12–19, d2 20–24, e1 16 –24, e2 53 –80, f2 57–82, h1 17–24, h2 266–353, h3 241–357, ps1 63–84. Ventral idiosoma. Epimerite I free. Seta c3 filiform, situated posterior to humeral shields. Epigynum almost semicircular, situated between tips of epimerites I. Oviporus long, shaped as an inverted Y, apodemes of oviporus extending to level of trochanters III (Figs. 7A, 10E). Anus close to posterior margin of idiosoma. Length of ventral setae: 1a 35–50, 3a 14–25, 4a 40–52, 4b 38, c3 20–23, ps2 33–49, ps3 17–22, g 20–29. Distance between setae: 1a:4b 64–78, 3a:4b 46–68, 4b:g 18–24, 4a:g 55–68, 4a:ps3 149–169. Legs (Figs. 9A–D, 10G, Table 1). Length of tarsi I–IV: 47–53, 60–63, 58–66 and 61–87, respectively. Length of tarsus I and II ca. 2 times of each greatest width. Setation of legs I–IV (excluding proral setae p and q): tarsi 8–8–4–6; tibiae 1–1–1–1; genua I–IV 2–2–0–0; femora I–IV 1–1–0–0; trochanters I–IV 0–1–1–0; coxae I–IV 1–0–1–0. Solenidiotaxy of legs I–II–III–IV: tarsi 2–1–0–0, tibiae 1–1–1–1, genua 1–1–1–0. Ambulacral discs I with smooth margin, discs of leg II–IV with ca. 10 denticles on distal edges. MALE. Dorsal idiosoma (Figs. 8A–B, 9E, 10C–D). Gnathosoma as in male, length 71–81, width 86–101. Body length including gnathosoma 425–450. Idiosoma transversally ovate, slightly wider than long, length 340–394 and greatest width 423–443, width including lateral membranes 437–448. Length of hysterosoma 281–297. Prodorsal shield as in female, length 75–92 and width 213–228. Setae vi spiculiform, situated near anterior end of prodorsal shield. Setae si narrowly lanceolate, and setae se long sword-shaped, separated by 94–105. Scapular shields represented by small bow-shaped sclerites. Lateral margins of hysterosoma with narrow membranes stretching from sejugal furrow to bases of setae f2. Hysteronotal shield: roughly ovate, wider than long, anterior margin slightly convex, length 245–273 and greatest width 388–401, most surface with numerous polygonal network pattern, area near posterior end additionally with small pits (Figs. 10C, D). Setae cp and c3 as in female. Setae d2, e2, f2, h2, h3, ps1, ps2 situated near lateral margins of hysterosoma. Setae d2 filiform. Setae e2, f2, ps2 and ps1 dilated, daggershaped; macrosetae h2 and h3 long dilated along all their length, gradually attenuate to apices. Distance between setae and hysteronotal gland openings: si:si 37–46, se:se 94–105, c1:d1 77–90, d1:gl 74–96, d2:gl 99–108, e1:gl 54–67, e1:h1 80–88, h1:h1 125–131, h2:h2 209–244, h3:h3 140–171, ps1:ps1 81–120. Length of dorsal setae: vi 13–18, si 95–107, se 148–165, c1 18–23, c2 69–91, d1 12–18, d2 22–30, e1 12 –15, e2 77–87, f2 61–74, h1 14–17, h2 186–342, h3 223–294, ps1 67–82. Ventral idiosoma. Epimerites I free, closer to each other than in female. Genital apparatus at level of trochanters IV. Aedeagus hook-shaped (Fig. 10F), 43–46 long. Both pairs of genital papillae and setae g approximately at midlength of genital arch. Anus close to posterior margin of opisthosoma. Adanal suckers situated at level of anus, disc-shaped, 23–26 in diameter, with radial patterns. Cupules ip close to f2. Length of ventral setae: 1a 57–74, 3a 43–53, 4a 26–40, 4b 28–49, c3 28–43, ps2 65–70, ps3 25–30, g 30–34. Legs (Fig. 9E, Table 1). Shape and setation as in female except tarsus IV. Length of tarsi I–IV 45–66, 45–65, 53–73, 61–81. Setae d and e of tarsus IV small spine-like, situated in distal 1/3 of segment. Remarks. Among four species of the genus Freyanopterolichus associated with ibises (Mégnin & Trouessart 1884; Trouessart & Mégnin 1885; Gaud & Mouchet 1959; Gaud 1982b), F. nipponiae is most close to F. chorioptoides (Mégnin & Trouessart, 1884) from Bostrychia carunculata (Threskiornithidae) in having: in both sexes, the idiosoma almost circular in shape, macrosetae h2 and h3 dilated, and idiosomal setae f2, and ps1 dagger-shaped; and in females, setae ps2 dagger-shaped. Freyanopterolichus nipponiae can be distinguished from the latter in the following combination of characters. In both sexes, the idiosoma is transversely ovate and slightly wider than long, tarsi I, II are approximately 2 times longer than greatest wide, macrosetae h2 and h3 are dilated along all their length and gradually attenuate to apices; in males, the terminal cleft is absent; in females, setae ps1 are as long as the distance between their bases. In both sexes of F. chorioptoides, the idiosoma is approximately as long as wide, tarsi I, II are approximately 3 times longer than their greatest wide, dilatations of macrosetae h2 and h3 are developed only in the basal part and have a noticeable angular extension on the inner margin; in males, the terminal cleft is shaped as a wide and shallow concavity; in females, setae ps1 are nearly two times as long as the distance between their bases., Published as part of Waki, Tsukasa & Shimano, Satoshi, 2022, Redescription of two parasitic feather mites sampled from the last two Crested Ibises, Nipponia nippon (Temminck, 1835) (Pelecaniformes: Threskiornithidae) lived in Japan, pp. 136-150 in Zootaxa 5116 (1) on pages 144-146, DOI: 10.11646/zootaxa.5116.1.7, http://zenodo.org/record/6364386, {"references":["Dubinin, V. B. (1953) Feather mites (Analgesoidea). Part II. Families Epidermoptidae and Freyanidae. Fauna SSSR, Paukoobraznye. Vol. 6. Fasc. 6. AN SSSR, Moscow-Leningrad, 411 pp. [in Russian]","Gaud, J. & Atyeo, W. T. (1996) Feather mites of the world (Acarina: Astigmata): The supraspecific taxa. Musee royal de l'Afrique centrale, Annales, Sciences zoologiques, 277 (Pt. 1 & 2), 1 - 193 (text) & 1 - 436 (illustrations).","Waki, T. & Shimano, S. (2020) A report of infection in the crested ibis Nipponia nippon with feather mites in current Japan. Journal of the Acarological Society of Japan, 29 (1), 1 - 8. https: // doi. org / 10.2300 / acari. 29.1","Kuroki, T., Tsurumi, M. & Nagahori, M. (2020) Comparison of species composition of feather mite Fauna on the Crested Ibis Nipponia nippon among populations from Japan, the Korean Peninsula, and Shaanxi Province, China (Reared in Japan). Yamashina Institute for Ornithology, 52, 113 - 123. [in Japanese with English summary] https: // doi. org / 10.3312 / jyio. 52.113","Megnin, P. & Trouessart, E. L. (1884) Diagnoses des especes et genres nouveaux de Sarcoptides plumicoles. Analgesinae, de la collection du Musee d'Angers. La Naturaliste, Serie 2, 6, 394 - 397.","Gaud, J. & Mouchet, J. (1959) Acariens plumicoles (Analgesoidea) parasites des oiseaux du Cameroun III. Dermoglyphidae. Annales de Parasitologie humaine et comparee, 34, 149 - 208. https: // doi. org / 10.1051 / parasite / 1959341149","Gaud, J. (1982 b) Acariens Sarcoptiformes plumicoles des oiseaux Ciconiidormes d'Afrique III. Parasites des Threskiornithidae. Revue de Zoologie Africaine, 96, 701 - 730."]}

Genus Freyanopterolichus Dubinin, 1953 Type species: Freyanopterolichus nipponiae Dubinin, 1953, by original designation. The genus Freyanopterolichus currently includes 10 species occurring on ibises (Threskiornithidae), storks (Ciconiidae), hamerkops (Scopidae) and cranes (Gruidae) (Trouessart & Mégnin, 1885; Dubinin 1953; Gaud & Mouchet, 1959; Gaud 1982a, 1982b). Within the family, this genus is clearly characterized by the following combination of characters. In both sexes, two internal vertical setae vi are present, epimerites I are free, the prodorsal shield is wide triangular and encompasses both pairs of scapular setae, the ambulacral discs of legs I are larger than those of legs II; in females, macrosetae h2 and h3 are long whip-like and usually dilated in basal part, the epigynum is situated between the extremities of epimerites I, and the oviporus is moved anterior and situated between levels of trochanters II and III., Published as part of Waki, Tsukasa & Shimano, Satoshi, 2022, Redescription of two parasitic feather mites sampled from the last two Crested Ibises, Nipponia nippon (Temminck, 1835) (Pelecaniformes: Threskiornithidae) lived in Japan, pp. 136-150 in Zootaxa 5116 (1) on page 144, DOI: 10.11646/zootaxa.5116.1.7, http://zenodo.org/record/6364386, {"references":["Dubinin, V. B. (1953) Feather mites (Analgesoidea). Part II. Families Epidermoptidae and Freyanidae. Fauna SSSR, Paukoobraznye. Vol. 6. Fasc. 6. AN SSSR, Moscow-Leningrad, 411 pp. [in Russian]","Gaud, J. & Mouchet, J. (1959) Acariens plumicoles (Analgesoidea) parasites des oiseaux du Cameroun III. Dermoglyphidae. Annales de Parasitologie humaine et comparee, 34, 149 - 208. https: // doi. org / 10.1051 / parasite / 1959341149","Gaud, J. (1982 a) Acariens Sarcoptiformes plumicoles des oiseaux Ciconiidormes d'Afrique II. Parasites des Threskiornithidae. Revue de Zoologie Africaine, 96, 335 - 357.","Gaud, J. (1982 b) Acariens Sarcoptiformes plumicoles des oiseaux Ciconiidormes d'Afrique III. Parasites des Threskiornithidae. Revue de Zoologie Africaine, 96, 701 - 730."]}

Compressalges nipponiae Dubinin, 1950 (Figs. 2–6) [Japanese name: Toki-umoudani] Compressalges nipponiae Dubinin, 1950: 537–540, fig. 1, 2; 1951: 29, fig. 16; 1956: 557–560, figs. 271–273; Gaud & Atyeo 1996: 79, fig. 216; Waki & Shimano 2020: 1–8, fig. 1; Kuroki et al. 2020, fig.1. Material examined: 4 females and 4 males (MPM Coll. No. 21820) from feathers of Nipponia nippon (Temminck, 1835) (Threskiornithidae), JAPAN, Niigata Prefecture, Sado Island, Ministry of the Environment, Sado Japanese Crested Ibis Conservation Center, 21, August 1993 –4, October 1994, feather coll. Dr. Yoshinori Kaneko. Description. FEMALE (Figs. 2A–B, 4A–D, 5A–B, 6A–B, D). Dorsal idiosoma. Gnathosoma: subcapitulum nearly square-shaped, length 97–126, greatest width 79–84. Body length including gnathosoma 506–558 long and 247–272 wide. Idiosoma 439–497 long, hysterosoma 349–377 long. Prodorsal shield strongly enlarged posteriorly, shaped as inverted mushroom, covering most part of prodorsum, anterior part with network pattern, posterior part with transverse striation, length 110–117, width of posterior part 117–124 (Figs. 2B, 5A). Setae ve rudimentary, situated on lateral margins of prodorsal shield, setae si and se near posterior margin of prodosal shield, setae se separated by 94–109. Hysteronotal shield enlarged in anterior part and gradually attenuate posteriorly, anterior margin with striations and shaped as trapezoid, close to prodorsal shield, length 329–374, greatest width 216–242, and surface with numerous small circular lacunae (Figs. 2B, 5B). Setae cp 153–187 long. Setae c2 thin spiculiform, 37–43 long. Setae c3 spiculiform, 34–39 long. Setae h3 spiculiform 78–94 long. Hysteronotal gland openings gl between levels of d2 and e2, cupules im between d2 and gl. Distances between setae and hysteronotal gland opening: se:se 87–92, si:si 51–55, d2: e2 56–76, d2:gl 31–44, e1:gl 63–75, h1: h1 23–29, h2:h2 57–63, h3:h3 45–52. Length of other dorsal setae: vi 8–11, se 94–109, c1 7–10, d1 9–13, d2 12–23, e1 13 –16, e2 11 –13, f2 17–22, h1 10–13, h2 210–252. Ventral idiosoma. Anus close to posterior margin of idiosoma. Oviporus shaped as an inverted Y, situated at level of sejugal furrow (Fig. 2A, 6A). Epigynum bow-shaped, 46–66 long and 41–62 wide. Cupules ih near lateral edges of opisthosoma at level of ps3. Length of ventral setae: 1a 38–56, 3a 49–52, 4a 47–57, 4b 20–28, ps2 10–11, ps3 9–13. Copulatory opening situated terminally, near posterior margin between setae ps1 (Fig. 2A, 6B). Legs (Figs. 4A–D, Table 1). Length of tarsi, I–IV: 49–57, 46–54, 50–56, 58–69. Setation of legs I–IV: tarsi 8–7–6–6; tibiae 1–1–1–1; genua 2–2–0–0; femora 1–1–0–0; trochanters 1–1–1–0; coxae I–IV 1–0–1–0. Solenidiotaxy of legs I–IV: tarsi 2–1–0–0, tibiae I 1–1–1–1, genua 2–1–1–0. Famulus epsilon ɛ short spine-like (Figs. 4A, 6D), close to solenidion ω1; setae ba of tarsi I, II distant from bases of solenidia ω1. MALE. (Figs. 3A–B, 4E, 5C–D, 6C). Dorsal idiosoma. Gnathosoma similar to that of female, length 88–101, width 66–83. Body length, including gnathosoma, 471–494, greatest width 230–253. Idiosoma length 400–442, hysterosoma length 316–329. Prodorsal shield as in female, length 103–111, width of posterior part 107–113. Setae si and se in posterior part of prodosal shield, setae se separated by 79–84. Hysteronotal shield strongly enlarged in anterior part and gradually attenuate posteriorly, anterior margin trapezoid-shaped, slightly concave in middle part, length 266–300, greatest width 186–212, surface with small circular lacunae in central area (Fig. 5C), and with large net-like ornamentation in posterior part (Fig. 5D). Setae c2 and c3 spiculiform. Opisthosomal lobes small, elongate, almost parallel-sided, length 42–47, greatest width 26–27. Terminal cleft U-shaped, greatest width 9–14. Outer margins of opisthosomal lobes with lateral membranes spreading from bases to posterior one third. Setae h1 narrowly lanceolate and situated on inner margins of opisthosomal lobes. Setae f2 spiculiform, situated near outer margins of opisthosomal lobes. Setae ps2 narrowly lanceolate, situated on lateral margins of opisthosomal lobes. Macrosetae h2 and h3 situated on posterolateral angles of opisthosomal lobes. Setae ps1 wide fan-shaped with radial striation, situated on terminus of opisthosomal lobes. Cupules im and hysteronotal gland openings gl between d2 and e2. Distances between setae and hysteronotal gland opening: se:se 79–86, si:si 46–54, d2: e2 77–94, d2:gl 59–76, e1: gl 30–38, h1:h1 11–15, h2:h2 60–68 h3:h3 45–52. Length of dorsal setae: vi 5–9, si 4–7, se 39–49, cp 141–205, c1 5–8, c2 28–38, c3 25–33, d1 5–8, d2 6–8, e1 4 –7, e2 6, f2 18–25, h1 22–25, h2 197–241, h3 183–229. Ventral idiosoma. Genital apparatus between trochanters IV, flanked laterally with setae g and ps3. Setae ps3 occasionally duplicated, additional pair smaller and situated anterior to original ps3 (Fig. 6C). Genital papillae posterior to setae 4a. Aedeagus stylet-like, 30–34 in length. Adanal suckers barrel-shaped, retracted in opisthosoma, 20–24 in length and 16–25 in width. Anus near anterior end of terminal cleft. Epimerites IVa long; adanal apodemes well developed, situated immediately posterior and parallel to epimerites IVa. Cupules ih near posterior ends of opisthosomal lobes. Length of ventral setae: 1a 32–39, 3a 31–42, 4a 28–38, 4b 23–28, ps1 31–34, ps2 23–29, ps3 13–17, g 21–27. Legs (Fig. 4E, Table 1): Legs IV hypertrophied. Length of tarsi I–IV: 46–51, 42–49, 45–50, 67–70. Leg setation as in female. Seta e of tarsus IV filiform, seta d small spiculiform and situated at level of distal 1/4 of this segment. Remarks. In the original description of Compressalges nipponiae and in the subsequent work, Dubinin (1950, 1956) erroneously interpreted the hysteronotal glands, which are dark-colored and have longitudinal striation, as some ventral setae modified in “large funnel-shaped suckers”. Gaud & Atyeo (1996) provided correct illustrations of this mite, but their drawing were too small-sized and did not show some tiny details of the genital apparatus, adanal suckers and opisthosoma of male., Published as part of Waki, Tsukasa & Shimano, Satoshi, 2022, Redescription of two parasitic feather mites sampled from the last two Crested Ibises, Nipponia nippon (Temminck, 1835) (Pelecaniformes: Threskiornithidae) lived in Japan, pp. 136-150 in Zootaxa 5116 (1) on pages 138-142, DOI: 10.11646/zootaxa.5116.1.7, http://zenodo.org/record/6364386, {"references":["Dubinin, V. B. (1950) Features of the structures of the fastening apparatus of the feather mite Compressalges nipponiae V. Dubinin, gen. and sp. nov. Proceedings of the USSR Academy of Sciences, 70, 537 - 540. [in Russian]","Gaud, J. & Atyeo, W. T. (1996) Feather mites of the world (Acarina: Astigmata): The supraspecific taxa. Musee royal de l'Afrique centrale, Annales, Sciences zoologiques, 277 (Pt. 1 & 2), 1 - 193 (text) & 1 - 436 (illustrations).","Waki, T. & Shimano, S. (2020) A report of infection in the crested ibis Nipponia nippon with feather mites in current Japan. Journal of the Acarological Society of Japan, 29 (1), 1 - 8. https: // doi. org / 10.2300 / acari. 29.1","Kuroki, T., Tsurumi, M. & Nagahori, M. (2020) Comparison of species composition of feather mite Fauna on the Crested Ibis Nipponia nippon among populations from Japan, the Korean Peninsula, and Shaanxi Province, China (Reared in Japan). Yamashina Institute for Ornithology, 52, 113 - 123. [in Japanese with English summary] https: // doi. org / 10.3312 / jyio. 52.113","Dubinin, V. B. (1956) Feather mites (Analgesoidea). Part III. Family Pterolichidae. Fauna SSSR, Paukoobraznye, Vol. 6, Fasc. 7. Moscow - Leningrad, publisher: AN SSSR, 814 pp. [in Russian]"]}

Waki, Tsukasa, Sasaki, Mizuki, Matsubara, Hajime, Shimano, Satoshi (2021): The First Report of the Feather Mite Pseudalloptinus milvulinus (Acariformes: Pterolichidae) from the Black Kite Milvus migrans in Japan. Species Diversity 26: 273-279, DOI: 10.12782/specdiv.26.273

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Yanagisawa, Shizuma, Sakamaki, Yositaka, Jantarit, Sopark, Shimano, Satoshi (2021): A New Species of the Genus Eucorydia (Blattodea: Corydiidae) from Chiang Mai in Northern Thailand. Species Diversity 26 (2): 191-195, DOI: 10.12782/specdiv.26.191, URL: http://dx.doi.org/10.12782/specdiv.26.191

Yanagisawa, Shizuma, Hiruta, Shimpei F., Sakamaki, Yositaka, Shimano, Satoshi (2021): A New Species of the Genus Eucorydia (Blattodea: Corydiidae) from the Miyako-jima Island in Southwest Japan. Species Diversity 26 (2): 145-151, DOI: 10.12782/specdiv.26.145, URL: http://dx.doi.org/10.12782/specdiv.26.145

Hygrobates (Lurchibates) incognitus sp. nov. Goldschmidt, Nishikawa & Shimano Material examined: Holotype female, slide mounted in glycerine jelly, preparation no. CIB INV 0023, parasitic on Paramesotriton guangxiensis (Amphibia, Caudata, Salamandridae); newt was collected in China, Guangxi Zhuang Autonomous Region, Ningming (mites were collected from unnumbered voucher specimens stored in the CIB collection, without detailed geographic information), preserved in 70% ethanol; mite was attached to the axilla and groin of the newt (CIB 200404064). Paratypes: Two females CIB INV 0024, same newt specimen than holotype; CIB INV 0025, same collecting data, different newt specimen (CIB 200404060). Distribution: All specimens of H. (L.) incognitus sp. nov. were collected from Paramesotriton guangxiensis. The new species is probably limited to the same distribution as its host (southern Guangxi, China and northeastern Cao Bang Province, Vietnam (Frost 2021)). Derivatio nominis: incognitus (Latin = unknown, not visible, unexpected); referring to the fact that this species in the molecular analysis (28S) could not be separated from macrochela sp. nov.. Diagnosis: Coxal field relatively broad; gnathosoma anterior heavily curved; female genital plates broad kidney-shaped, flanking posterior 4/5 of genital opening; P-4 relatively slender, proximo-ventral extension of P-5 large, blunt cone-shaped; cheliceral claw relatively curved, slender. Description, Male: Unknown. Description, Female (n = 3): Idiosoma rounded-oval, L/W ratio 1.43 (1.40), L/W 1680 (1344)/1176 (960); fused anterior coxae of both sides elongated, triangular, Cx-I + II L/W 432 (360–400)/636 (564–582), ratio 0.68 (0.62–0.71), medio-posterior margin only slightly extended by secondary sclerotization, posteriorly convex, lateral extension straight, pointing postero-laterally, Cx-I basal width 228 (209–212); gnathosoma anteriorly heavily curved, broad, rounded tips far projecting, lateral margin with Cx-I posteriorly clearly converging (Fig. 30); anterior and posterior coxal groups laterally clearly separated, medially diverging; posterior coxae far separated, inclined heart-shaped, antero-medial margin curved, medial edges nose-shaped projecting, formed by Cx-IV only, Cx-IV nearly rectangular, posterior margin transverse, lateral margin convex (Fig. 30), posterior coxal groups (Cx-III + Cx-IV of one side) L/W 354 (318–322)/341 (294–306); genital field overall broad oval, acetabular plates broad kidney-shaped, slightly inclined to lateral, flanking 4/5 of genital opening, pre- and postgenitale laying completely under integument, genital field L/W 258 (234–252)/378 (336–342), single genital plate L/W 174 (178–188)/102 (92–94); acetabula of similar size, irregular oval, Ac-2 and Ac-3 laying beside each other, L/W Ac-1 76 (79–82)/43 (41–42), Ac-2 82 (82–85)/46 (41–48), Ac-3 89 (83–84)/40 (49–51), 18/20 (20/18, 20/22) setae of similar size on anterior, lateral and posterior margin of each plate, antero-medial of the plates 4/3 (2/3, 3/4), postero-medial 3/0 (0, 1/1) additional setae in the soft integument beside the pre- and postgenital sclerites (see Fig. 30); all legs slender, bearing many heavy setae, especially dorso-distally at basal segments (Figs. 31–35); measurements (L/H) of distal leg segments: I-leg-5 325 (322–325)/54 (54), I-leg-6 266 (240–258)/50 (49–50); II-leg-5 342 (342–346)/54 (54– 55), II-leg-6 276 (259–270)/54 (52–54); III-leg-5 378 (372–384)/56 (58–60), III-leg-6 312 (300–312)/54 (54–55); IV-leg-3 236 (236–242)/72 (67–68), IV-leg-5 396 (396–414)/58 (54), IV-leg-6 354 (342–355)/52 (53); chelicerae strong; cheliceral claws very large, curved, distally sharply pointed, dorsal margin in the distal half with strong serration continuing proximally in a lateral serration, medially and laterally striated (Figs. 36, 37); palps strong, relatively slender (Figs. 38–40), ventral margin of P-2 very slightly concave (sharp bend in Fig. 38 probably misshapen), antero-ventral corner with single denticles (in some specimens none), P-3 ventrally straight to slightly convex with loose field of denticles in distal 2/3, P-4 relatively long and slender, slightly curved, with a pair of ventral setae in the distal fourth; P-5 with a blunt, cone-shaped ventro-proximal projection, dorso-distally with a compact, denticle-like distal claw, distally with a pair of large, strong, similar, ventrally-buckled claws; mouthpart measurements: Chelicera H 151 (basal part in all specimens broken), claw L 261, curvation of cheliceral claw 26°; palp total L 617 (604–616), L/H P-1 71 (61–63)/87 (85–87), P-2 161 (165–169)/99 (96–101), P-3 122 (118–122)/87 (87–92), P-4 212 (204–212)/76 (63–68), P-5 52 (52–54)/45 (54–56).

Annotations to the subgenus Lurchibates Comparison Hygrobates (Lurchibates) to Hygrobates (s. str.) The most obvious character separating Lurchibates from other subgenera of Hygrobates, beside the life-style as newt parasites, are the by far larger, stronger and overall more massive mouthparts (Figs. 46, 47). In contrast, the morphology of the idiosoma and the legs does not show any specific modifications (Figs. 45 vs. 48). Sexual dimorphism in Lurchibates As mentioned before (Goldschmidt & Fu 2011, Goldschmidt et al. 2020), sexual dimorphism in the subgenus Lurchibates mainly refers to the shape of the coxal field (the basal width of Cx-I in general is larger in females), thus the main difference is found in the shape of the genital field (following the general pattern of Hygrobates) (Figs. 51 vs. 52): In males the acetabular plates are fused forming a single wing-like, dumbbell- or apple-shaped genital plate (Fig. 51); in females the acetabular plates are always separated, not connected with the pre- and post-genital sclerites, flanking the genital opening (in some species shifted to posterior), their shape varies from kidney- to crescent-shaped (Fig. 52). Furthermore females are generally larger than males and show a wider variation in size: E.g. the idiosoma length in H. (L.) aloisii in the type series varies from 948–1134 in males and 1488–1932 in females (in the present study from 756–852 in males, 864–1320 in females). In H. (L.) macrochela sp. nov. the idiosoma length varies from 660–924 in males and 882–1536 in females. The size differences of males vs. females are again intensified by an enormous increase of size in ovigerous females: H. (L.) aloisii (present study), 864–1260 (non-ovigerous females) vs. 1320 (ovigerous female); H. (L.) macrochela sp. nov., 882–1104 (non- ovigerous females) vs. 1284–1536 (ovigerous females) (Figs. 52, 53). The so far studied specimens of most species included as well several ovigerous females, however the number of eggs per specimen is rather variable within the species. Based on the current data no differences between the species are visible in this respect. The number of eggs per ovigerous female varies overall between one and 100 (average 38, median 42). Whereas the egg size not varies much, neither between nor among species (overall diameter 140–165). The sexual dimorphism documented in H. (L.) intermedius, idiosoma L 900 (males) vs. 1824 (female), is probably strongly biased by the fact, that the only female used in the study was an ovigerous specimen (Goldschmidt et al. 2020). Additional information to the so far known species of Lurchibates So far the only records of the subgenus Lurchibates have been the species descriptions. Consequently any information on the morphological variability was exclusively based upon the type series of the respective species. In the present study (Goldschmidt et al. 2020, present paper), besides the seven new species, further specimens of three out of four already known species were found: H. (L.) forcipifer, H. (L.) ancistrophorus and H. (L.) aloisii. Therefore, we are providing some additional measurement data for these species (see Tab. 2, appendix): The new measurement data for the males of H. (L.) forcipifer and the single female of H. (L.) ancistrophorus are mostly within the already known range, just the male genital field of H. (L.) forcipifer is slightly smaller in the present study (Tab. 2, appendix). Yet the measurements of the three males and four females of H. (L.) aloisii are greatly extending the known size range of that species, the specimens of the present study (males as well as females) are clearly smaller than the ones of the type series (Tab. 2, appendix). Distribution of the so far known species of Lurchibates The genus Lurchibates seems to be limited to SE-Asia, the distribution of the individual species could be derived from the distribution pattern of their respective hosts (Fig. 54). Relationship among Lurchibates species based on morphometric analysis In order to confirm and visualize the separation of the so far known species of Hygrobates (Lurchibates), and to get an idea of phylogenetic relationships among the species, a character matrix was analyzed. Principal Component Analysis (PCA) based on raw data results in an ordination of individuals showing most species grouped together (Fig. 55) whereas there are as well certain overlaps between all species when the first two principal axes are plotted, except for ancistrophorus which is placed distant from all others. The first three components account for 79.2% of total variation (PC1 58.1, PC2 15.1, PC3 6.0). Highest loadings on PC1 showed the variables W of Cx-III and Cx-IV (one side) with 0.221 and P1-dorsal length with 0.296. On PC 2 the variables Ac-1 width, Ac-2 width and Ac-3 width showed the highest loadings with 0.367, 0.394 and 0.482 respectively. The variables Ac-2 length, P-5 length and P-2 height showed the highest loadings on PC3 with values of 0.366, 0.394 and 0.378. Variation explained by PC1 is related to size and shows that size contributes considerably to species separation. PCA on size corrected data results in larger overlaps between the species, whereas individuals are scattered on two distinctly separated “groups”: The first group with macrochela, robustipalpis, intermedius and malosimilis and the second group including ancistrophorus, pilosus, aloisii, incognitus and forcipifer. Lurchibates salamandrarum is clearly isolated in this analysis, by far separated from remaining individuals (Fig. 56). The first three components account for 74.1% of total variation (PC1 40.1, PC2 20.8, PC3 13.2). Highest loadings were shown in the variable width of coxa I+II. Non metric Multidimensional Scaling (NMDS) on raw data shows a clear morphometric distinction between all species, again with certain overlaps but with a stress of 0.1004 (Fig. 57)., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F., Pfingstl, Tobias, Jiang, Jian-Ping & Shimano, Satoshi, 2021, Systematics, distribution and morphology of the newt parasitic water mites of the subgenus Lurchibates Goldschmidt & Fu, 2011 (Acari, Hydrachnidia Hygrobatidae, Hygrobates Koch, 1837), including the description of four new species and a key to all so far known species, pp. 1-36 in Zootaxa 4985 (1) on pages 15-21, DOI: 10.11646/zootaxa.4985.1.1, http://zenodo.org/record/4929035, {"references":["Goldschmidt, T. & Fu, V. (2011) Description of Hygrobates aloisii sp. nov. from Hong Kong, a new species Hygrobates (Lurchibates) subgen. nov. (Acari, Hydrachnidia, Hygrobatidae), with data on the parasite - host relationship to the Hong Kong Newt Paramesotriton hongkongensis (Amphibia, Caudata, Salamandridae). Zoologischer Anzeiger, 250, 19 - 31. https: // doi. org / 10.1016 / j. jcz. 2010.10.002","Goldschmidt, T., Nishikawa, K., Hiruta, S. F. & Shimano, S. (2020) Description of three new water mite species of Hygrobates Koch, 1837 (Lurchibates Goldschmidt & Fu, 2011) (Acari, Hydrachnidia, Hygrobatidae), parasitic in newts of the genera Paramesotriton and Pachytriton (Amphibia, Caudata, Salamandridae) from China. Zootaxa, 4768 (1), 25 - 42. https: // doi. org / 10.11646 / zootaxa. 4768.1.3","Goldschmidt, T., Gerecke, R. & Alberti, G. (2002) Hygrobates salamandrarum sp. nov. (Acari, Hydrachnidia, Hygrobatidae) from China: the First Record of a Freshwater Mite Parasitizing Newts (Amphibia, Urodela). Zoologischer Anzeiger, 241, 297 - 304. https: // doi. org / 10.1078 / 0044 - 5231 - 00073","Chang, M. L. Y. (1933) On the salamanders of Chekiang. Contributions from the Biological Laboratory of the Science Society of China. Zoological Series, 9, 305 - 328.","Goldschmidt, T. & Koehler, G. (2007) New species of the Hygrobates salamandrarum - group (Acari, Hydrachnidia, Hygrobatidae) from SE-Asia. Zoologischer Anzeiger, 246, 73 - 89. https: // doi. org / 10.1016 / j. jcz. 2007.01.001","Stuart, B. L. & Papenfuss, T. J. (2002) A new salamander of the genus Paramesotriton (Caudata: Salamandridae) from Laos. Journal of Herpetology, 36, 145 - 148. https: // doi. org / 10.1670 / 0022 - 1511 (2002) 036 [0145: ANSOTG] 2.0. CO; 2","Bourret, R. (1934) Notes herpetologiques sur l'Indochine francaise. VI. Sur diverses collections de serpents appartenants a l'Universite de Hanoi. VII. Une salamandre nouvelle vivant au Tonkin. Annexe au Bulletin General de l'Instruction Publique. Hanoi, 1934, 83 - 84.","Huang, Z. - Y., Tang, Z. - Y. & Tang, Z. - M. (1983) A new species of the genus Trituroides from Guangxi, China. Acta Herpetologica Sinica / Liangqi baxing dongwu yanjiu. New Series. Chengdu, 2 (2), 37 - 39.","Myers, G. S. & Leviton, A. E. (1962) The Hong Kong newt described as a new species. Occasional Papers. Division Of Systematic Biology, Stanford University, 10, 1 - 4.","Yuan, Z. - Y., Zhao, H. - P., Jiang, K., Hou, M., He, L., Murphy, R. W. & Che, J. (2014) Phylogenetic relationships of the genus Paramesotriton (Caudata: Salamandridae) with the description of a new species from Qixiling Nature Reserve, Jiangxi, southeastern China and a key to the species. Asian Herpetological Research, 5, 67 - 79. https: // doi. org / 10.3724 / SP. J. 1245.2014.00067","Wu, Y. - K., Jiang, K. & Hanken, J. (2010) A new species of newt of the genus Paramesotriton (Salamandridae) from southwestern Guangdong, China, with a new northern record of P. longliensis from western Hubei. Zootaxa, 2494 (1), 45 - 58. https: // doi. org / 10.11646 / zootaxa. 2494.1.3","Nishikawa, K., Jiang, J. - P., Matsui, M. & Mo, Y. - M. (2011) Unmasking Pachytriton labiatus (Amphibia: Urodela: Salamandridae), with description of a new species of Pachytriton from Guangxi, China. Zoological Science, 28 (6), 453 - 461. https: // doi. org / 10.2108 / zsj. 28.453"]}

Hygrobates (Lurchibates) malosimilis sp. nov. Goldschmidt, Nishikawa & Shimano Material examined: Holotype male, slide mounted in glycerine jelly, preparation no. CIB INV 0022, parasitic on Pachytriton inexpectatus (Amphibia, Caudata, Salamandridae); newt was collected in China, Guangxi Zhuang Autonomous Region, Jinxiu Yao Autonomous County, Mt. Dayao 24°06’27’’ N, 110°13’52’’ E, 1210 m a.s.l. on September 10 th 2008, preserved in 70% ethanol; mite was attached to the hindlimb of the newt (CIB GX20081008). Distribution: The only specimen of H. (L.) malosimilis sp. nov. was collected from Pachytriton inexpectatus from Guangxi Zhuang Autonomous Region, China. The new species is at maximum limited to the same distribution as its host, which is so far known from Eastern Guizhou, southwestern and southern Hunan, extreme northwestern Guangdong, and northern and eastern Guangxi, China. Derivatio nominis: malosimilis; derived from malum (Latin = apple) and similis (Latin = similar); named for the apple-shaped male genital field. Diagnosis (only male): Idiosoma relatively small; anterior coxal group narrow, very slender, basally of intermediate width (compared to most other species of the subgenus), regularly rounded; posterior coxal group medially pointed, very irregular triangular; male genital field apple-shaped; palp relatively compact, P-3 ventro-distally with a patch of denticles, P-4 relatively short, slightly curved, proximo-ventral extension of P-5 blunt, flat cone-shaped, large distal claws of different size (medial smaller), smaller one slender; cheliceral claw curved, extended in distal half. Description, Male (n = 1): Idiosoma rounded-oval, L/W ratio 1.23, L/W 684/558; fused anterior coxae of both sides slender, narrow triangular, Cx-I + II L/W 264/324, ratio 0.81, secondary sclerotization at medio-posterior margin rather small (posteriorly forming a regular curve, lateral apodemes oriented towards antero-lateral); Cx-I basal width 139, gnathosoma slender, posteriorly only slightly and regularly converging, lateral separation posteriorly only reaching slightly more than half the length of anterior coxal group, widely fused with the posterior part of the first coxae, anterior tip of gnathosoma slightly projecting, anterior margin straight to concave (Fig. 20); posterior coxal groups (Cx-III + Cx-IV of one side) L/W 222/174, anterior and posterior coxal groups laterally very close, medially diverging; posterior coxae irregularly triangular, medial edges pointed, formed by Cx-IV only, anterior and posterior margin heavily undulating, lateral margin slightly undulating, nearly parallel to longitudinal axis (Fig. 20); genital field apple-shaped, anteriorly nose-like pointed, posteriorly with broad, deep, apically rounded indentation, with 17 pairs of setae irregularly arranged at outer margin and laterally beneath elongate-oval genital opening (Fig. 20), genital field L/W 162/210; Ac-1 irregularly drop-shaped, L/W 63/31, Ac-2 lenticular, L/W 77/32, Ac-3 irregularly rectangular drop-shaped, L/W 70/42; genital skeleton partly distorted, L/W 207/150 as far as visible brachia distalia strong, brachia proximalia strong, parallel to longitudinal axis (Fig. 21); all legs slender (especially segments 4–6), bearing several heavy setae, mainly distally at most segments (Figs. 22–25); measurements (L/H) of distal leg segments: I-leg-5 186/40, I-leg-6 158/38; II-leg-5 198/42, II-leg-6 180/42; III-leg-5 216/43, III-leg-6 200/42; IV-leg-3 148/50, IV-leg-5, 228)/41, IV-leg-6 212/40; chelicerae very strong, with a relatively short, high basal segment and mid-sized basal groove; cheliceral claws very heavy, strongly curved, sharply pointed, dorsal margin in the distal half extended, with strong serration, medially and laterally striated (Figs. 26, 27); palps relatively compact (Figs. 28, 29), P-2 dorsally regularly curved, ventral margin slightly concave, without denticles, ventro-distal corner of P- 3 rounded, distal half with small field of denticles, P-4 relatively short, slightly curved, with a pair of ventral setae near ventro-distal corner, distal margin convex, rotated ventrally, P-5 with a well developed, blunt cone-shaped ventro-distal projection, dorso-distal seta relatively slender, distal claws clearly different, medial one slender, slightly curved, lateral one clearly stronger, nearly rectangularly hooked; mouthpart measurements: Chelicera total L 342, H 104, L/H ratio 3.3, basal segment L 254, claw L 160, basal segment/claw ratio 1.6; curvation of cheliceral claw 30°, palp total L 376, L/H P-1 33/61, P-2 110/82, P-3 73/67, P-4 118/49, P-5 42/38. Female (unknown), Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F., Pfingstl, Tobias, Jiang, Jian-Ping & Shimano, Satoshi, 2021, Systematics, distribution and morphology of the newt parasitic water mites of the subgenus Lurchibates Goldschmidt & Fu, 2011 (Acari, Hydrachnidia Hygrobatidae, Hygrobates Koch, 1837), including the description of four new species and a key to all so far known species, pp. 1-36 in Zootaxa 4985 (1) on pages 8-10, DOI: 10.11646/zootaxa.4985.1.1, http://zenodo.org/record/4929035

Hygrobates (Lurchibates) fragmentarius sp. nov. Goldschmidt, Nishikawa & Shimano Unfortunately the only specimen of this new species is severely damaged, nevertheless the few morphological data as well as the molecular information (28S) clearly demonstrate that it’s a new species. Even though, a complete species description is not possible, we are giving all available information here. Material examined: Fragment of unknown sex, slide mounted in glycerine jelly, preparation no. CIB INV 0026, parasitic on Paramesotriton yunwuensis (Amphibia, Caudata, Salamandridae); newt was collected in China, Guangdong (mite was collected from unnumbered voucher specimen stored in the CIB collection, without detailed geographic information) preserved in 70% ethanol; mite was attached to the groin of the newt. Distribution: The only specimen of H. fragmentarius sp. nov. was collected from Paramesotriton yunwuensis. The new species is probably limited to the same distribution as its host, southwestern Guangdong Province, China (Wu et al., 2010). Derivatio nominis: fragmentarius (Latin = fragmented, fragmentary); referring to the poor condition of the only available specimen. Diagnosis: Anterior coxal group very slender; gnathosoma anterior heavily curved; palp strong and slender; P-4 relatively slender, proximo-ventral extension of P-5 large, blunt cone-shaped; cheliceral claw relatively curved. Description, Sex unknown (n = 1): Idiosoma heavily damaged; fused anterior coxae of both sides elongated, triangular, Cx-I + II L/W 366/492, ratio 0.74, medio-posterior margin clearly extended by secondary sclerotization, posteriorly narrow rounded, postero-lateral extension slightly hook-shaped, Cx-I basal width 122, ratio Cx-I L/Cx-I basal width 3.0; gnathosoma anteriorly heavily curved, rounded tips far projecting, lateral margin with Cx-I posteriorly clearly converging, connection with posterior part of Cx-I narrow; gnathosoma and Cx-I anteriorly separated by deep indentation (Fig. 41); chelicerae strong; cheliceral claws very large, curved, distally sharply pointed, dorsal margin in the distal half with strong serration that continues proximally in a lateral serration, medially and laterally striated, basal part with long ventral groove (Figs. 42, 43); palps strong, relatively slender (Fig. 44), ventral margin of P-2 straight, without denticles, P-3 ventrally straight with field of denticles in distal 2/3, P-4 relatively long and slender, slightly curved, with a pair of ventral setae in the distal fourth; P-5 with a blunt, cone-shaped ventro-proximal projection, dorso-distally with a compact, denticle-like distal claw, distally with a pair of large, strong, similar, ventrally curved claws; mouthpart measurements: Chelicera L 442, H 122, basal segment L 287, claw L 202, curvation of cheliceral claw 26°; palp total L 556, L/H P-1 52/69, P-2 148 /96, P-3 110/63, P-4 194/61, P-5 52/54. Remarks on the new species described above: Hygrobates (Lurchibates) macrochela sp. nov. and H. (L.) malosimilis sp. nov. are very similar in the shape of the palp and especially the very characteristic heavy, distally thickened cheliceral claw, separating them from all other Lurchibates species (Figs. 7–10, 16–19, 26–29). They are clearly separated from each other in the shape of the male genital field: Wider in H. (L.) macrochela (Fig. 1), rather narrow in H. (L.) malosimilis (Fig. 20); and the shape of the anterior coxal group: Slightly more slender in H. (L.) macrochela, wider in H. (L.) malosimilis (Cx-I L/basal W 1.5 –1.8 in H. (L.) macrochela, 1.9 in H. (L.) malosimilis; Cx-I+II L/ W 0.67 –0.75 in H. (L.) macrochela, 0.81 in H. (L.) malosimilis). The narrow anterior coxal group (Fig. 20) is as well separating H. (L.) malosimilis from all other species of the subgenus (Cx-I+II L/ W 0.81 in H. (L.) malosimilis, 0.62–0.77 in all other species). Hygrobates (Lurchibates) incognitus sp. nov. unfortunately so far is only known in the female, nevertheless the species can clearly be separated from the other known species of Lurchibates by the shape of their mouthparts and genital field: H. (L.) incognitus is bearing a relatively slender, “normal” palp (P-4 L/H 2.8–3.3), whereas H. (L.) salamandrarum (P-4 L/H 2.1) and H. (L.) robustipalpis (P-4 L/H 2.5–2.6) are characterized by rather compact palps; H. (L.) ancistrophorus has a unique palp with P-5 missing the ventro-distal cone which is present in all other species of the subgenus. H. (L.) incognitus has large and strongly curved cheliceral claws, but these are not distally thickened, as they are in H. (L.) macrochela and H. (L.) malosimilis; Furthermore the cheliceral claw is relatively straight (claw curve 19–21°) in H. (L.) aloisii, H. (L.) intermedius and H. (L.) forcipifer, as well as in H. (L.) macrochela and H. (L.) malosimilis, whereas the cheliceral claw is more curved (claw curve 26–27°) in H. (L.) incognitus, H. (L.) pilosus (Figs. 36, 37) and H. (L.) fragmentarius sp. nov. (Figs. 42, 43). Moreover, in H. (L.) incognitus the acetabular plates are rather kidney-shaped, small, posteriorly not extending beyond the post-genital sclerite, the arrangement of the acetabula is rather triangular (Ac-3 beside Ac-2), there are three to four setae laying free in the integument antero-medial to the acetabular plates (Fig. 30); in contrast, in females of H. (L.) macrochela the crescent-shaped acetabular plates are slightly extending beyond the post-genital sclerite, the acetabula are forming an arc (Ac-3 clearly posterior to Ac-2), there are no setae antero-medial to the acetabular plates (Fig. 11); in H. (L.) pilosus and H. (L.) forcipifer the acetabular plates are as well not reaching beyond the post-genital sclerite, but in these species the acetabula are rather forming an arc; in H. (L.) aloisii and H. (L.) intermedius the acetabular plates are – as in H. (L.) incognitus – rather kidney-shaped, small, but they are shifted posteriorly, at least reaching the posterior end of the post-genital sclerite. Even though the only specimen of Hygrobates (Lurchibates) fragmentarius sp. nov. is heavily broken and fragmented, the species is clearly separated from all other so far known species of the subgenus by very long and slender gnathosoma and anterior coxae (Fig. 41), especially Cx-I is basally very narrow (Cx-I L/basal W 3.0 vs. 1.5–2.8 in other species); the palps are slender, especially P-3 (L/H 1.8 vs. 0.9–1.4 in all other species)., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F., Pfingstl, Tobias, Jiang, Jian-Ping & Shimano, Satoshi, 2021, Systematics, distribution and morphology of the newt parasitic water mites of the subgenus Lurchibates Goldschmidt & Fu, 2011 (Acari, Hydrachnidia Hygrobatidae, Hygrobates Koch, 1837), including the description of four new species and a key to all so far known species, pp. 1-36 in Zootaxa 4985 (1) on pages 14-15, DOI: 10.11646/zootaxa.4985.1.1, http://zenodo.org/record/4929035, {"references":["Wu, Y. - K., Jiang, K. & Hanken, J. (2010) A new species of newt of the genus Paramesotriton (Salamandridae) from southwestern Guangdong, China, with a new northern record of P. longliensis from western Hubei. Zootaxa, 2494 (1), 45 - 58. https: // doi. org / 10.11646 / zootaxa. 2494.1.3"]}

Subgenus Hygrobates (Lurchibates) Goldschmidt & Fu, 2011 Diagnosis: Integument fine, irregularly striated, glandularia little sclerotized, lateral eyes not in capsules; genital field with 3 pairs of acetabula; excretory pore with very weak sclerotization; legs without swimming hairs, I-leg-5 ventro-distally with a pair of strong setae, I-leg-6 straight; gnathosoma large, posteriorly broadly fused to the anterocoxal plate; cheliceral claws extremely large, with dorsal margin distally curved and serrated, proximally concave; palpus robust, P-2 without projection, P-3 ventrally smooth or denticulate, P-5 short, compact and formed like a grasping organ due to the presence of a proximo-ventral extension (flat to extended, pointed) and two strong, curved distal claws. There is a clear sexual dimorphism in the shape of the genital field (for details see Goldschmidt & Fu 2011)., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F., Pfingstl, Tobias, Jiang, Jian-Ping & Shimano, Satoshi, 2021, Systematics, distribution and morphology of the newt parasitic water mites of the subgenus Lurchibates Goldschmidt & Fu, 2011 (Acari, Hydrachnidia Hygrobatidae, Hygrobates Koch, 1837), including the description of four new species and a key to all so far known species, pp. 1-36 in Zootaxa 4985 (1) on page 3, DOI: 10.11646/zootaxa.4985.1.1, http://zenodo.org/record/4929035, {"references":["Goldschmidt, T. & Fu, V. (2011) Description of Hygrobates aloisii sp. nov. from Hong Kong, a new species Hygrobates (Lurchibates) subgen. nov. (Acari, Hydrachnidia, Hygrobatidae), with data on the parasite - host relationship to the Hong Kong Newt Paramesotriton hongkongensis (Amphibia, Caudata, Salamandridae). Zoologischer Anzeiger, 250, 19 - 31. https: // doi. org / 10.1016 / j. jcz. 2010.10.002"]}

Scolopendra alcyona Tsukamoto & Shimano, sp. nov. [New Japanese name for Scolopendra alcyona sp. nov.: Ry��jin-��mukade (���������������); new English name for Scolopendra alcyona sp. nov.: Halcyon giant centipede] (Figs 3B, C, 4���9) Material examined. Holotype: Male (NSMT-My 511, TS-20180914-01), Oku, Kunigami-son, Kunigami-gun, Okinawa-jima Island, Okinawa prefecture, Japan (26��49���04.6���N, 128��16���52.3���E), 14 Sep. 2018, coll. Hutoshi Taira. Paratypes: sex unknown (NSMT-My 512, TS-20180912-01), Yona, Kunigami-son, Kunigami-gun, Okinawajima Island, Okinawa prefecture, Japan, 12 Sep. 2018, coll. Katsuya Tsukamoto; sex unknown (RUMF-ZD-02001, TS-20180301-01), Yona, Kunigami-son, Kunigami-gun, Okinawa-jima Island, Okinawa prefecture, Japan, 1 Mar. 2018, coll. Takeshi Sasaki; 2 sex unknown (NSMT-My 513, TS-20180522-01; NSMT-My 514, TS-20180522-02), Nakachi, Kumejima-cho, Shimajiri-gun, Kume-jima Island, Okinawa prefecture, Japan, 22 May 2018, coll. Humiyasu Sato. All type specimens except TS-20180301-01 are deposited at the Collection of Myriapoda, Department of Zoology, NSMT, and TS-20180301-01 is deposited at RUMF. Non-type specimens: sex unknown (NSMT-My 515, TS-2018MMDD-01), Nuha, Ogimi-son, Kunigami-gun, Okinawa-jima Island, Okinawa prefecture, Japan, 2018, coll. Zento Touyama; Female (NTU TS-19690814-01), Hengchun, Pingtung City, Taiwan, 14 Aug. 1969, coll. Mitsuo Takano. TS-2018MMDD-01 is deposited at the Collection of Myriapoda, Department of Zoology, NSMT. Taiwanese specimen (TS-19690814-01) are deposited at NTU. Etymology. This epithet ���alcyona��� derived from the name of a Greek mythological figure, ���Alcyone���. In one version of the story, Alcyone was turned into a common kingfisher (halcyon bird) by Zeus (Gresseth 1964). This epithet is a metaphor for halcyon bird-like jade green legs seen in the Okinawa-jima Island population, and its amphibious behavior. Diagnosis. 17���20 antennal articles, basal 6 articles glabrous; cephalic plate with small puncta; each tooth-plate with 4���9 teeth; tergites (4, 5) 6���20 with complete paramedian sutures; tergite of ultimate leg-bearing segment without median suture; sternites 2���19 with well-defined, incomplete paramedian suture; coxopleural process with 2 apical spines and 1 subapical spine; ultimate legs long and slender with ratio of width and length of prefemur 1:3, ratio of width and length of femur 1:4 and ratios of lengths of prefemur and femur 1.06:1; femur and tibia 1.13:1; tibia and tarsus 1 1.33:1; tarsus 1 and tarsus 2 2.15:1; dorsally flattened ultimate leg prefemora with 2 VL, 1 M, 2 DM and 1���3 SP; tarsal spurs on legs 1���19; gonopods absent, penis present. Scolopendra alcyona sp. nov. is similar to all other congeners in East and Southeast Asia (see ���Morphological comparison among Japanese and Taiwanese Scolopendra ���, and Tables 6 and 7). S. alcyona sp. nov. can be distinguished from S. japonica by the absence gonopods in the male, and incomplete paramedian sutures. S. alcyona sp. nov. can be distinguished from S. multidens by the ratio of the length of femur and tibia, and tibia and tarsus (femur and tibia 1.1:1; tibia and tarsus 1 1.3: 1 in S. alcyona sp. nov.; femur and tibia 1.7:1, tibia and tarsus 1 1.8: 1 in S. multidens). S. alcyona sp. nov. can be distinguished from S. dawydoffi by the tergite number on which the complete paramedian sutures start (T4���T 6 in S. alcyona sp. nov.; T2 or T 3 in S. dawydoffi). S. alcyona sp. nov. can be distinguished from S. mutilans and S. subspinipes by the absence gonopods in the male, and the presence of a tarsal spur only on legs 1���19. Holotype description (variation of paratypes is given in parenthesis). Body length in 100% Ethanol 130 mm rounding off (TS-20180301-01: 190 mm, TS-20180912-01: 180 mm, TS-20180522-01: 140 mm, TS-20180522-02: 130 mm). Antenna (Figs 4A, 7A, 9A, C) with 18 articles (17���20 article in paratypes); basal 6 articles glabrous. Cephalic plate (Figs 4A, 7A, 9C) with small puncta entirely; short anterior median sulcus; paramedian suture absent. Article 2 of second maxillary telopodite with spur. Second maxillary pretarsus 0.36 times as long as article 3 of second maxillary telopodite, rounded at the tip, with dorsal and ventral spurs. Coxosternite (Figs 4B���D, 7B���D) surface smooth, without median suture; tooth-plates longer than wide, with 7 + 7 teeth in two groups (8 + 7 in TS-20180301-01, 9 + 8 in TS-20180912-01, 5 + 4 in TS-20180522-01 and 5 + 7 in TS-20180522-02) and with transverse basal sulci. Forcipule (Figs 4B���D, 7B) surface smooth; process of trochanteroprefemur with teeth in two groups, 1 apical tooth and 1(3) inner teeth. Tergite (Figs 4E, 7E) surface smooth. Anterior margin of T1 overlapped by cephalic plate. Complete paramedian sutures on TT6���20 (paramedian sutures starting on TT 4���6 in paratypes); margination on TT6���20 (margination starting on TT 6���11 in paratypes). Tergite of ultimate leg-bearing segment (Figs 5A, 8A) without median suture or depression; posterior margin of tergite of ultimate leg-bearing segment curved posteriorly; ratio of width and length of tergite of ultimate leg-bearing segment 1:1. Sternite (Figs 4F, 7F) surface smooth; SS2���19 with paramedian sutures, posteriorly incomplete, extending to approximately 65���76% length of sternite. Sternite of ultimate leg-bearing segment (Figs 5B, 8B) with lateral margins converging posteriorly and posterior margin convex posteriorly; surface with shallow median longitudinal depression. Coxopleuron (Figs 5B, C, 8B, C) with numerous pores, pore-field extending into coxopleural process covering most of its surface, except for a narrow pore-free area, dorsal margin of pore-field gradually elevated anteriorly. Coxopleural process moderately long, with 2 apical spines and 1 subapical spine (0���2 apical and 0���2 subapical spines in paratypes), without lateral spine; pore-free area of coxopleural process extending ventrally 79���93% length from distal part of coxopleural process to posterior margin of sternite of ultimate leg-bearing segment (73���100% in paratypes). All legs without setae and tibial spur. One ventrodistal tarsal spur on tarsus 1 of legs 1���19. Ultimate legs (Figs 6, 8A, B, 9D) long and slender with ratio of width and length of prefemur 1:3, ratio of width and length of femur 1:4 and ratios of lengths of prefemur and femur 1.06:1; femur and tibia 1.13:1; tibia and tarsus 1 1.33:1; tarsus 1 and tarsus 2 2.15:1. Prefemur of ultimate leg flattened dorsally with acute blackish spines row as below: 2 VL, 1 M, 2 DM and 2���3 SP (1���3 SP in paratypes). In the male holotype, and a female non-type specimen (TS-19690814-01), genital segments (Figs 5D, E) everted and projected beyond distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly. Gonopods absent. Sternite of genital segment 2 well developed in male. Tergite of genital segment small, horseshoeshaped and lacking setae. Lamina subanalis between genitalia and anal valves; lamina adanalis between anal valves and tergite of genital segment. Penis present. Coloration in living condition as in Figs 9A���D. Brownish black on cephalic plate and T1; greenish black on remaining tergites; bluish black on antenna; light brown on coxosternite and forcipule (but forcipular tarsungulum blackish); pale green on all sternites; bluish black on eupleuria, with greenish black integuments; orange on coxopleura; basal part of legs 1���20 yellow, distal part of legs 1���20 and ultimate legs greenish blue (all legs yellow in Kume-jima Island specimens); pale blue on intermediate sternite. Distribution: Ryukyu Archipelago (Okinawa-jima Island, Kume-jima Island) and Taiwan. S. alcyona sp. nov. specimens examined in this study were collected in Okinawa-jima Island, Kume-jima Island, and Taiwan, but S. alcyona sp. nov. could be also distributed in Tokashiki-jima Island, Ishigaki-jima Island, and Iriomote-jima Island (Masashi Sugimoto and Taku Shimada, personal communication). Habitat: Scolopendra alcyona sp. nov. lives along streams that flow through the forest. It is often found under stones nearby the streams., Published as part of Tsukamoto, Sho, Hiruta, Shimpei F., Eguchi, Katsuyuki, Liao, Jhih-Rong & Shimano, Satoshi, 2021, A new amphibious species of the genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) from the Ryukyu Archipelago and Taiwan, pp. 465-494 in Zootaxa 4952 (3) on pages 483-489, DOI: 10.11646/zootaxa.4952.3.3, http://zenodo.org/record/4690551, {"references":["Gresseth, G. K. (1964) The myth of alcyone. Transactions and Proceedings of the American Philological Association, 95, 88 - 98. https: // doi. org / 10.2307 / 283785"]}

Preliminary taxonomic key to the Japanese-Taiwanese Scolopendra species In the following key, body coloration is used for discriminating S. mutilans and S. subspinipes because of the lack of the other discriminative morphological characters. However, the reliability of body coloration in discrimination of Scolopendra species is controversial (Chao & Chang 2003, Shinohara et al. 2015). Further morphological surveys using adequate specimens of both these species are needed. 1 T21 with median suture.......................................................... S. morsitans Linnaeus, 1758 - T21 without median suture.............................................................................. 2 2 Gonopods present in male.............................................................................. 3 - Gonopods absent in male............................................................................... 5 3 Tarsal spur only on legs 1���19.......................................................... S. japonica Koch, 1878 - Tarsal spur on legs 1���19(20)............................................................................ 4 4 Head and body dark brown in adult (monochromatic).................................... S. subspinipes Leach, 1816 - Head and T1 red or reddish brown, body dark green in adult (dichromatic)...................... S. mutilans Koch, 1878 5 Ratio of length of prefemur and femur 1.2:1, femur and tibia 1.7:1, tibia and tarsus 1 1.8:1; tarsus 1 and tarsus 2 1.5:1; cephalic plate with large puncta; penis absent in male........................................... S. multidens Newport, 1844 - Ratio of length of prefemur and femur 1.1:1; femur and tibia 1.1:1; tibia and tarsus 1 1.3:1; tarsus 1 and tarsus 2 2.9:1; cephalic plate with small puncta; penis present in male............................. S. alcyona Tsukamoto & Shimano, sp. nov., Published as part of Tsukamoto, Sho, Hiruta, Shimpei F., Eguchi, Katsuyuki, Liao, Jhih-Rong & Shimano, Satoshi, 2021, A new amphibious species of the genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) from the Ryukyu Archipelago and Taiwan, pp. 465-494 in Zootaxa 4952 (3) on page 490, DOI: 10.11646/zootaxa.4952.3.3, http://zenodo.org/record/4690551, {"references":["Chao, J. L. & Chang, H. W. (2003) The scolopendromorph centipedes (Chilopoda) of Taiwan. African Invertebrates, 44 (1), 1 - 11.","Shinohara, K., Takano, M. & Ishii, K. (2015) Chilopoda. In: Aoki, J. (Ed.), Pictorial keys to soil animals of Japan. Tokay University Press, Tokyo, pp. 873 - 910.","Linnaeus, C. (1758) Systema naturae. Editio Decima. Salvius, Holmiae, 824 pp.","Koch, L. (1878) Japanesische Arachniden und Myriopoden. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 27, 285 - 295.","Leach, W. E. (1816) A tabular view of the external characters of four classes of animals, which Linne arranged under Insecta; with the distribution of the genera composing three of these classes into orders, & c. and descriptions of several new genera and species. Transactions of the Linnean Society of London, 11, 306 - 400. https: // doi. org / 10.1111 / j. 1096 - 3642.1813. tb 00065. x","Newport, G. (1844) XIII. - A list of the species of Myriapoda, order Chilopoda, contained in the cabinets of the British Museum, with synoptic descriptions of forty-seven new species. The Annals and Magazine of Natural History, including Zoology, Botany and Geology, Series 1, 13 (82), 94 - 101. https: // doi. org / 10.1080 / 03745484409442576"]}

Pfingstl, Tobias, Hiruta, Shimpei F., Nemoto, Takamasa, Hagino, Wataru, Shimano, Satoshi (2021): Ameronothrus twitter sp. nov. (Acari, Oribatida) a New Coastal Species of Oribatid Mite from Japan. Species Diversity 26 (1): 93-99, DOI: 10.12782/specdiv.26.93, URL: http://dx.doi.org/10.12782/specdiv.26.93

Pyatakovestia boninensis Morino and Miyamoto, 2015 is reported based on single female specimen from Ototojima Island, the Bonin Islands, Japan. This is the first record of the species in Ototojima Island. The specimen referred to P. boninensis differs from the original description (type locality: Hahajima Island) by the shape of coxal gill of pereopod 6 and the number of marginal setae of outer ramus of uropod 2., 小笠原諸島弟島からハマトビムシ科のオガサワラホソハマトビムシを初めて記録した。弟島から得られた雌1個体について分類学的検討を行った結果、原記載とは第6胸肢の底節鰓の形状、および第2尾肢外肢縁棘数について変異がみられることが分かった。

The Ameronothridae are recorded for the first time from Japanese coasts with the new species Ameronothrus yoichi sp. n. from Hokkaido. The report of this species represents the most southern occurrence of an Ameronothrus species in the Asian Pacific region. Ameronothrus yoichi sp. n. can be easily distinguished from its congeners by the conspicuously pusticulate body surface and the loss of dorsal companion setae d on all genua in the adult stage. Based on adult and juvenile morphology, a close relation to Ameronothrus maculatus and A. schneideri is suggested. Ameronothrus yoichi sp. n. is classified as a lichenivorous inhabitant of sediment-free rocky coastal substrates. Due to a lack of genetic sequence data of nearly all ameronothrid species a molecular genetic comparison is yet unfeasible, but a Bayesian inference tree based on the 18S rRNA gene shows a paraphyletic clustering of the ameronothrid A. yoichi sp. n. and Paraquanothrus grahami. http://www.zoobank.org/urn:lsid:zoobank.org:pub:5B772E2C-7D5E-4C86-9955-AB84A84C50DA, Graphical abstract

Pfingstl et al. 2021. Ameronothrus twitter sp. nov. (Acari, Oribatida) a New Coastal Species of Oribatid Mite from Japan. Species Diversity.

Supplement 1. old

Hygrobates salamandrarum Goldschmidt, Gerecke and Alberti, 2002 (Acari, Hydrachnidia, Hygrobatidae) is the firstknown water mite parasitizing adult newts. After its description, three further species of newt-parasitizing mites were described and the subgenus Lurchibates Goldschmidt and Fu, 2011 was proposed for the group, now containing four newtparasitizing mites. Until now, each water mite species parasitizes a different newt species (Table 1), suggesting possible mite-newt co-speciation. In order to test this hypothesis, we need an accurate taxonomy of both of the hosts and parasites. However, the taxonomy of those Asian newt genera known to be parasitized by Lurchibates mites has been substantially revised after the initial description of the H. salamandrarum. To account for these taxonomic changes, we here revise the host-parasite species list and amend the host species name as shown in Table 1. Hygrobates salamandrarum was collected from the body of Pachytriton labiatus (Unterstein, 1930) obtained via pet trade (Goldschmidt et al., 2002). However the identity of this newt species was revised by Nishikawa et al. (2011a). These authors found that the type specimen of Pac. labiatus belonged to a different genus (Paramesotriton), and its name therefore changed to Paramesotriton labiatus (Unterstein, 1930). By this taxonomic change, the Pachytriton newt that had been considered as ��� Pachytriton labiatus ��� and from which the mites were collected, came to have no name. Further, Nishikawa et al. (2011a) found that the ��� Pachytriton labiatus ��� could be separated into two species, and consequently they revived Pachytriton granulosus Chang, 1933 for populations from Zhejiang and Anhui provinces, and newly described Pac. inexpectatus Nishikawa, Jiang, Matsui and Mo, 2011 for those from Guizhou, Henan, and Guangdong provinces and Guangxi Zhuang Autonomous Region. After that, Nishikawa et al. (2011b) revised the taxonomy of Pachytriton, and the Anhui population of Pac. granulosus was described as Pac. feii Nishikawa, Jiang, and Matsui, 2011 and some populations from Guangxi, sympatric to Pac. inexpectatus, were described as Pac. moi Nishikawa, Jiang, and Matsui, 2011. Unfortunately, the symbiotypes (host newt specimens) of H. salamandrarum were not assigned and the specimens were lost. The specimens originated from the pet-trade, and therefore no locality information is available for them. Judging from the measurements and color pattern from the photos of the newts (Fig. 1), the host most probably can be assigned to be Pac. granulosus Chang, 1933. At least one of the specimens (Fig. 1A and C) is a matured male with swollen cloaca. Its snout-vent length [SVL] can be estimated from the available photos, which contained a scale, as ca. 61mm. This adult male size is within the ranges of P. granulosus (range 59.0��� 78.9mm, mean 69.0mm) and P. feii (range 58.5���90.5mm, mean 73.3mm: Nishikawa et al., 2009) but smaller than other species of the genus (Nishikawa et al., 2011b). Further, the host newts had body proportions more similar to P. granulosus than to P. feii in the ratio of forelimb length versus SVL (male 22.2% and unknown sex 22.6% vs. male mean 23.2% in P. granulosus, but vs. male mean 25.8% and female mean 23.7% in P. feii) and that of hindlimb length versus SVL (male 24.4% and unknown sex 26.2% vs. male mean 25.3% in P. granulosus, but vs. male mean 29.6% and female mean 28.3% in P. feii [Nishikawa et al., 2009]). The color pattern is also similar between the host newts and P. granulosus (ventral markings are large and widely occupy the venter and cloacal marking connects with that of ventral tail: see Figs. 4 and 5 in Nishikawa et al., 2009), but not P. feii (markings are small and blotched and cloacal marking does not connect with that of ventral tail: see Fig. 5 in Nishikawa et al., 2011b). We, thus, propose to revise the symbiotypic species of H. salamandrarum as to be Pac. granulosus. For Hygrobates ancistrophorus Goldschmidt and Koehler, 2007, the mites were collected from Paramesotriton laoensis Stuart and Papenfuss, 2002, which has subsequently been transferred to a different genus, Laotriton Dubois and Raffa��lli, 2009 (Frost, 2019). Thus, the host newt of H. ancistrophorus is now Laotriton laoensis (Stuart and Papenfuss, 2002)., Published as part of Nishikawa, Kanto, Goldschmidt, Tom, Hiruta, Shimpei F. & Shimano, Satoshi, 2020, Taxonomic amendments of Southeast Asian newt species of the genera Pachytriton, Paramesotriton and Laotriton (Amphibia, Urodela, Salamandridae) parasitized by water mites of the subgenus Lurchibates (Hydrachnidia, Hygrobatidae, Hygrobates), pp. 297-300 in Zootaxa 4768 (2) on pages 297-299, DOI: 10.11646/zootaxa.4768.2.11, http://zenodo.org/record/3779798, {"references":["Goldschmidt, T., Gerecke, R. & Alberti, G. (2002) Hygrobates salamandrarum sp. nov. (Acari, Hydrachnidia, Hygrobatidae) from China: the first record of a freshwater mite parasitizing newts (Amphibia, Urodela). Zoologischer Anzeiger, 241, 297 - 304. https: // doi. org / 10.1078 / 0044 - 5231 - 00073","Goldschmidt, T. & Fu, V. (2011) Description of Hygrobates aloisii sp. nov. from Hong Kong, a new species Hygrobates (Lurchibates) subgen. nov. (Acari, Hydrachnidia, Hygrobatidae), with data on the parasite - host relationship to the Hong Kong Newt Paramesotriton hongkongensis (Amphibia, Caudata, Salamandridae). Zoologischer Anzeiger, 250, 19 - 31. https: // doi. org / 10.1016 / j. jcz. 2010.10.002","Unterstein, W. (1930) Beitrage zur Lurch- und Kriechtierfauna Kwangsi's. 2. Schwanzlurche. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 313 - 315.","Goldschmidt, T. & Koehler, G. (2007) New species of the Hygrobates salamandrarum - group (Acari, Hydrachnidia, Hygrobatidae) from SE-Asia. Zoologischer Anzeiger, 246, 73 - 89. https: // doi. org / 10.1016 / j. jcz. 2007.01.001","Stuart, B. L. & Papenfuss, T. J. (2002) A new salamander of the genus Paramesotriton (Caudata: Salamandridae) from Laos. Journal of Herpetology, 36, 145 - 148. https: // doi. org / 10.1670 / 0022 - 1511 (2002) 036 [0145: ANSOTG] 2.0. CO; 2","Myers, G. S. & Leviton, A. E. (1962) The Hong Kong newt described as a new species. Occasional Papers. Division Of Systematic Biology, Stanford University, 10, 1 - 4.","Nishikawa, K., Jiang, J. - P., Matsui, M. & Mo, Y. - M. (2011 a) Unmasking Pachytriton labiatus (Amphibia: Urodela: Salamandridae), with description of a new species of Pachytriton from Guangxi, China. Z o o l o g i c a l S c i e n c e, 28, 453 - 461. https: // doi. org / 10.2108 / zsj. 28.453","Chang, M. L. Y. (1933) On the salamanders of Chekiang. Contributions from the Biological Laboratory of the Science Society of China. Zoological Series, 9, 305 - 328.","Nishikawa, K., Jiang, J. - P. & Matsui, M. (2011 b) Two new species of Pachytriton from Anhui and Guangxi, China (Amphibia: Urodela: Salamandridae). Current Herpetology, 30, 15 - 31. https: // doi. org / 10.5358 / hsj. 30.15","Nishikawa, K., Jiang, J. - P., Matsui, M. & Chen, C. - S. (2009) Morphological variation in Pachytriton labiatus and a re-assessment of the taxonomic status of P. granulosus (Amphibia: Urodela: Salamandridae). Current Herpetology, 28, 49 - 64. https: // doi. org / 10.3105 / 018.028.0202","Dubois, A. & Raffaelli, J. (2009) A new ergotaxonomy of the family Salamandridae Goldfuss, 1820 (Amphibia, Urodela). Alytes, 26, 1 - 85."]}

Subgenus Hygrobates (Lurchibates) Goldschmidt & Fu, 2011 The new material treated in the present publication allows us to give a slightly modified and extended diagnosis of the subgenus compared with the original one given by Goldschmidt & Fu (2011). Diagnosis: Integument fine, irregularly striated, dorsal and postero-ventral muscle attachment sites not sclerotized; lateral eyes not in capsules; glandularia weakly sclerotized, gland pores and their accompanying setae closely associated, but not on a common platelet; genital field with three pairs of acetabula on more or less triangular plates, males with a single genital plate formed by the fusion of both acetabular plates with the strongly sclerotized pre- and postgenitale; excretory pore with very weak sclerotization; legs without swimming hairs, their claws with ventral claw blade, I-leg-5 ventro-distally with a pair of strong, short, blunt setae, distal leg segments straight; gnathosoma large, posteriorly broadly fused to the little extended antero-coxal plate, rostrum moderately projecting; cheliceral claws extremely large, with dorsal margin distally curved and serrated, proximally concave; palp robust, P-2 with- out projection, P-2 and P-3 ventrally smooth or denticulate, P-5 short, compact and formed like a grasping organ due to the presence of a proximo-ventral extension (flat to extended, pointed) and a pair of large, strong, curved distal claws; a fine distally ramified seta inserted ventro-laterally between the proximal projection and the strong palp claws; clear sexual dimorphism in the shape of the genital field, in general slight sexual dimorphism in the shape of the coxal field; no sexual dimorphism in the morphology of legs, palps and chelicerae., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F. & Shimano, Satoshi, 2020, Description of three new water mite species of Hygrobates Koch, 1837 (Lurchibates Goldschmidt & Fu, 2011) (Acari, Hydrachnidia, Hygrobatidae), parasitic in newts of the genera Paramesotriton and Pachytriton (Amphibia, Caudata, Salamandridae) from China, pp. 25-42 in Zootaxa 4768 (1) on page 26, DOI: 10.11646/zootaxa.4768.1.3, http://zenodo.org/record/3777804, {"references":["Goldschmidt, T. & Fu, V. (2011) Description of Hygrobates aloisii sp. nov. from Hong Kong, a new species of Hygrobates (Lurchibates) subgen. nov. (Acari, Hydrachnidia, Hygrobatidae), with data on the parasite - host relationship to the Hong Kong Newt Paramesotriton hongkongensis (Amphibia, Caudata, Salamandridae). Zoologischer Anzeiger, 250, 19 - 31. https: // doi. org / 10.1016 / j. jcz. 2010.10.002"]}

Hygrobates intermedius sp. nov. Goldschmidt, Nishikawa & Shimano Material examined: Holotype male, slide mounted in glycerine jelly, preparation no. CIB INV 0001, parasitic on Paramesotriton qixilingensis (Amphibia, Caudata, Salamandridae): newt was collected in China, Jiangxi Province, Yongxin County, Shenyuan Station; N 26�� 46��� 25.2������, E 114�� 10��� 53.7������, 375m a.s.l. on March 26th 2014, preserved in 70% Ethanol; mite was attached to the fore- or hindlimb of the newt (CIB 140328). Paratypes: one female CIB INV 0002 from fore- or hindlimb of the newt (CIB 140328), one male CIB INV 0003 from mouth cavity, limbs, or lateral body of the newt (CIB 140327, 29���31, or 33)���newts data are the same as holotype���s host. Distribution: All three specimens of H. intermedius sp. nov. have been recorded on Paramesotriton qixilingensis. The type locality of the newt (Qixiling Nature Reserve, Mount Shenyuan, Yongxin County, Jiangxi, China) is the only known collecting site for both, the newt and the mite. Derivatio nominis: intermedius (lat.)���located between, intermediate; named for the fact that most characters of the new species are somehow intermediate between all other species of Lurchibates. Diagnosis: Characters of the subgenus Lurchibates; anterior coxal group relatively narrow posteriorly (compared to most other species of the subgenus), posterior coxal group triangular; male genital field rounded dumbbell-shaped, regularly surrounded by many setae; female genital plates crescent-shaped, relatively short (flanking only posterior 2/3 of genital opening); palp relatively long and slender, P-3 ventrally with dense field of denticles, P-4 long, straight, proximo-ventral extension of P-5 blunt, cone-shaped; cheliceral claw relatively straight, long and slender. Description, Male (n = 2): Idiosoma rounded-oval, L/W ratio 1.34 (1.25), L/W 900 (900)/672 (720); fused anterior coxae of both sides slightly elongate (Cx-I + II L/W 294 (310)/462 (439)), medio-posterior margin straight to slightly convex, laterally extended by secondary sclerotization (swallow-tail shaped, lateral tips reaching under medial margin of Cx-III/IV), Cx-I basal W 160 (150); gnathosoma broad, posteriorly only slightly converging, widely fused with the posterior part of the first coxae, anterior margin of gnathosoma curved (Fig. 1); posterior coxal groups (Cx-III + IV of one side) L/W 288 (283)/270 (270), laterally closely approached to anterior plates (Cx- II postero-laterally overlapping antero-lateral corners of Cx-III), medially clearly diverging from anterior coxae, nearly regularly triangular in shape, small, rounded medial edges formed by Cx-IV only, Cx-IV short and broad, nearly triangular, posterior margin straight to slightly convex, medially pointing towards anterior, lateral margin straight to convex (Fig. 1); genital field dumbbell-shaped (Ac-1 and Ac-3 anteriorly and especially posteriorly extending beyond pre- and postgenitale), outline broad-oval (anteriorly and laterally rounded, posteriorly straight), pregenitale nose-shaped, anteriorly extended, genital opening relatively small, elongated egg-shaped, genital field L/W 198 (204)/ 348 (384); Ac-1 elongated oval, Ac-2 irregularly oval, Ac-3 elongated triangular, anteriorly pointed (L/W Ac-1, 104 (100)/54 (51), Ac-2, 118 (107)/51 (54), Ac-3, 113 (116)/77 (64)), 25���28 setae on each side of plate, arranged at the margins of the genital plate (Fig. 1); excretory pore slit-shaped; genital skeleton L/W 223 (252)/183 (181), compact (L/W ratio 1.22 (1.39)), shape typical for the subgenus, brachia distalia long, strong, laterally strongly curved, brachia proximalia strong, distally broadened, parallel to longitudinal axis, (Fig. 2); all legs slender, bearing many heavy setae (Figs. 3���6); measurements (L/H) of distal leg segments: I-leg-5, 252 (248)/54 (48), I-leg-6, 216 (199)/48 (50); II-leg-5, 264 (277)/54 (54), II-leg-6, 228 (240)/48 (49); III-leg-5, 294 (259)/56 (53), III-leg-6, 252 (222)/52 (48); IV-leg-3, 180 (175)/68 (66), IV-leg-5, 300 (306)/49 (54), IV-leg-6, 282 (270)/43 (48); chelicerae very strong, with a relatively short, high basal segment and long basal groove; cheliceral claws very large, slender, sharply pointed, dorsal margin in the distal third with strong serration that continues proximally in a lateral serration, medially and laterally striated (Figs. 7, 8); palps strong (Figs. 9, 10), ventral margin of P-2 slightly concave, antero-ventral margin rounded, without denticles, P-3 ventrally straight with a field of pointed denticles in the distal half, occasionally the distal two thirds, P-4 relatively long and straight, distally slightly curved, with a pair of ventral setae in the distal third; P-5 with a well developed, blunt cone-shaped ventro-proximal projection, dorsodistally P-5 bears one compact, denticle-like distal claw and ventro-distally a pair of similar large, strong, pointed, slightly curved claws; mouthpart measurements: chelicera total L 408 (394), maximum H 102 (96), L/H ratio 4.0 (4.1), basal segment L 259 (244), claw L 198 (190), basal segment/claw ratio 1.3 (1.3); curvation of cheliceral claw 21�� (20��), palp total L 486 (493), L/H P-1, 47 (45)/63 (68), P-2, 129 (134)/81 (80), P-3, 94 (94)/75 (75), P-4, 169 (173)/56 (56), P-5, 47 (47)/45 (47). Description, Female (n = 1): Idiosoma similar to male, much larger, L/W 1824/1464, and more rounded (L/W ratio 1.25); unpaired anterior coxal group slightly more compact, basally broader than in males (Cx-I + II L/W 408/589), basal width Cx-I 211; medio- and latero-posterior margin extended by secondary sclerotization, Cx-I latero-posteriorly with apodemes similar to those in males, however more hook-shaped; gnathosoma broadly fused with the posterior part of the first coxae, lateral margins straight, nearly parallel, coxal groups further separated than in males, medially heavily diverging (Fig. 11); paired posterior coxal groups well distanced from each other and from the anterior coxal group, L/W 356/320, roughly triangular in shape with medial edges only slightly diverging from longitudinal axis, postero-medial corner less pointed, medial margin formed by Cx-IV and Cx-III (Fig. 11); genital field far distant from coxae, rather small, with strongly sclerotized pre- and postgenitale, genital plates well distant from each other, separated by soft integument, slightly kidney-shaped, anteriorly rounded, flanking posterior 2/3 of genital opening, total genital field L/W 312/420; acetabula irregularly-oval (L/W Ac-1, 83/54, Ac- 2, 109/54, Ac-3, 101/58); 20/22 setae arranged mainly at the anterior, posterior and lateral margins of the genital plates, two, respectively three, setae anterior to genital plates (Fig. 11); legs similar to males, slightly more slender (Figs. 12���15); measurements of distal leg segments: L/H I-leg-5, 276/54, I-leg-6, 276/54; II-leg-5, 294/54, II-leg-6, 257/54; III-leg-5, 324/60, III-leg-6, 282/53; IV-leg-3, 228/85, IV-leg-5, 354/60, IV-leg-6, 312/48; anterior margin of the gnathosoma by far more projecting than in males, anterior half completely separated from Cx-I (Fig. 11); chelicerae similar to males, slightly more compact (Figs. 16, 17); palps similar to males (Figs. 18, 19); mouthpart measurements: chelicera total L 456, maximum H 126, L/H ratio 3.6, basal segment L 282, claw L 221, basal segment/claw ratio 1.3; cheliceral claw curve 21��; palp total L 550, L/H P-1, 54/78, P-2, 150/94, P-3, 110/85, P-4, 188/62, P-5, 47/49., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F. & Shimano, Satoshi, 2020, Description of three new water mite species of Hygrobates Koch, 1837 (Lurchibates Goldschmidt & Fu, 2011) (Acari, Hydrachnidia, Hygrobatidae), parasitic in newts of the genera Paramesotriton and Pachytriton (Amphibia, Caudata, Salamandridae) from China, pp. 25-42 in Zootaxa 4768 (1) on pages 27-31, DOI: 10.11646/zootaxa.4768.1.3, http://zenodo.org/record/3777804

Hygrobates pilosus sp. nov. Goldschmidt, Nishikawa & Shimano Material examined: Holotype male, slide mounted in glycerine jelly, preparation no. CIB INV 0008, parasitic on Paramesotriton yunwuensis (Amphibia, Caudata, Salamandridae): newt from pet shop, preserved in 99% Ethanol; mite was attached to the waist of the newt (KUHE 49342). Paratypes: three females CIB INV 0009, CIB INV 0010, CIB INV 0011, five males CIB INV 0012, CIB INV 0013, CIB INV 0014, CIB INV 0015, CIB INV 0016 from head, lateral body, or groin of the newt (KUHE 49340)���same data as holotype, but fixed by freezing (-80��C) [therefore the mites were in poor conditions and could only partly be measured and illustrated]. Distribution: All specimens of H. pilosus sp. nov. have been collected from Paramesotriton yunwuensis. The new species is probably limited to the same distribution as its host, which was described from Yunwu Mountains, Luoding, Guangdong Province, China, and is only known from there. Derivatio nominis: pilosus lat. hairy���referring to the many long setae at the margin of the male genital plate. Diagnosis: Characters of the subgenus; coxal field relatively slender (compared to other species of the subgenus), anterior coxal group triangular, posteriorly narrow, elongated, posterior margin rounded; male genital field broad-oval, bearing many long setae; female genital plates crescent-shaped, anteriorly narrow, rounded; P-4 relatively slender, proximo-ventral extension of P-5 large, triangularly pointed; cheliceral claw relatively straight, long and slender. Description, Male (n = 6): Idiosoma rounded-oval, L/W ratio 1.42 (1.36���1.39), L/W 888 (816���864)/624 (588���636); fused anterior coxae of both sides elongated, triangular (Cx-I + II L/W 312 (330���336)/442 (456���492)), medio-posterior margin extended by secondary sclerotization (posteriorly convex, curved tips of lateral extension reaching under medial margin of Cx-III), Cx-I basal width 136 (120���136); gnathosoma relatively long and slender, anteriorly only very slightly projecting, lateral margin with Cx-I posteriorly clearly converging, at fusion with the posterior part of the Cx-I relatively narrow (Fig. 39); anterior and posterior coxal groups laterally very close (but not overlapping), medially slightly diverging (the clear lateral separation at one side given in Fig. 39, is an artifact, as the integument of the respective side is broken between Cx-II and -III); posterior coxae very close to each other, triangular, medial edges smoothly rounded, formed by Cx-IV only, Cx-IV nearly rectangular, posterior margin transverse, slightly undulating, lateral margin straight, parallel to longitudinal axis of idiosoma (Fig. 39), posterior coxal groups (Cx-III + Cx-IV of one side) L/W 276 (270���288)/241 (269���288); genital plate very broad oval to wing-like, acetabular plates smoothly fused with strongly sclerotized pre- and postgenitale, genital opening broad egg-shaped, anteriorly obtuse-angled, genital field L/W 174 (198���216)/366 (354���390); acetabula relatively far distant from each other (about the width of an acetabulum), Ac-1 and -2 irregularly oval, Ac-3 triangularly rounded, L/W Ac-1 76 (63���82)/33 (36���40), Ac-2 70 (63���72)/43 (36���46), Ac-3 63 (61���80)/55 (57���63), 38���47 setae on each side of the plate (see Fig. 39) mainly arranged at the margins, lateral setae longer than anterior and posterior ones (Fig. 39); genital skeleton typical for the subgenus, large, L/W 277 (287���306)/181 (193���202) and slender, L/W ratio 1.53 (1.49���1.51), brachia distalia strong, laterally curved, oblique antero-medially to postero-laterally, brachia proximalia strong, distally broadened, parallel to longitudinal axis (left side slightly distorted in the illustrated specimen) (Fig. 40); all legs slender, bearing many heavy setae (Figs. 41���44); measurements (L/H) of distal leg segments: I-leg-5, 264 (264)/52 (48), I-leg-6, 218 (216)/54 (48); II-leg-5, 276 (276)/54 (54), II-leg-6, 234 (228)/54 (54); III-leg-5, 306 (318)/56 (56), III-leg-6, 252 (258)/54 (48); IV-leg-3, 174 (174)/72 (70), IV-leg-5, 324 (342)/54 (53), IV-leg-6, 294 (313)/52 (42); chelicerae strong, relatively slender; cheliceral claws very large, distally straight, sharply pointed, dorsal margin in the distal third with strong serration that continues proximally in a lateral serration, medially and laterally striated (Figs. 45, 46); palps strong (Figs. 47, 48), ventral margin of P-2 straight, anteroventral corner with very few denticles, P-3 ventrally straight with few denticles in distal half, P-4 relatively long, straight, distally slightly curved, with a pair of ventral setae in the distal third; P-5 with a well developed, coneshaped ventro-proximal projection, dorso-distally with a compact, denticle-like distal claw and ventro-distally a pair of large, strong, ventrally-directed curved claws similar to each other; mouthpart measurements: chelicera total L 420 (414���456), maximum H 102 (110���115), L/H ratio 4.1 (3.7���4.0), basal segment L 257 (240���273), claw L 197 (180���197), basal segment/claw ratio 1.3 (1.4���1.5); curvation of cheliceral claw 21��, palp total L 536 (530���543), L/H P-1, 52 (49���52)/68 (69���75), P-2, 146 (139���146)/94 (87), P-3, 103 (103���118)/82 (80���82), P-4, 186 (181���188)/63 (59���61), P-5, 49 (47���51)/49 (49���52). Description, Female (n = 3): Idiosoma similar to male, much larger, L/W 1800���1908 /1536���1440, and more rounded (L/W ratio 1.17���1.33); unpaired anterior coxal group wider and relatively shorter than in males, L/W 390���438/576���601, medio- and latero-posterior margin extended by secondary sclerotization as in males, but lateroposterior apodemes of Cx-I not reaching under Cx-III, basal width Cx-I 208���237; gnathosoma relatively wider and shorter than in males, lateral margins posteriorly clearly converging, broadly fused with the posterior part of the first coxae (Fig. 49); paired posterior coxal groups more distanced from each other and from the anterior coxal group than in males, roughly triangular in shape with medial edges formed by Cx-IV only, medial margin undulating, Cx-IV trapezoid, L/W 354���360/336���342 (Fig. 49); genital field rounded (exact position of strongly sclerotized pre- and postgenitale not clear as all females in poor conditions due to fixation technique, see above), genital plates slender kidney-shaped, total genital field L/W 186/318; acetabula irregularly-oval to triangular (Fig. 49), L/W Ac-1, 70���75/33���39, Ac-2, 86���89/34���45, Ac-3, 75���96/51���57; 21���24 setae arranged mainly at the anterior, posterior and lateral margins of the genital plates, none to one setae anterior to genital plates (Fig. 49); measurements of distal leg segments: L/H I-leg-5, 319/-, I-leg-6, 264/-; II-leg-5, 336���366/54���60, II-leg-6, 282���288/54; III-leg-5, 402���410/55��� 66, III-leg-6, 330���348/54; IV-leg-3, 252���258/72, IV-leg-5, 376���438/56���66, IV-leg-6, 336���378/54���48 (legs not illustrated due to poor conditions); chelicerae similar to males, slightly stronger (Figs. 50, 51); palps similar to males (Fig. 52); mouthpart measurements: chelicera total L 541���600, maximum H 144���156, L/H ratio 3.5���3.9, basal segment L 354���362, claw L 271���275, basal segment/claw ratio 1.3; cheliceral claw curve 20���22��; palp total L 632���658, L/H P-1, 61���68/89���108, P-2, 172���181/108, P-3, 118���125/90���99, P-4, 221���230/71���73, P-5, 56/61���63. Remarks: The new species Hygrobates intermedius sp. nov. from Jiangxi Province, China is most similar to H. robustipalpis sp. nov. ���males of both species are characterized by a slightly dumbbell-shaped genital field. However, in H. intermedius sp. nov. the outline of the male genital field is rather regularly oval, whereas in H. robustipalpis sp. nov. the outline is apple-shaped. Furthermore, in the latter the anterior coxal group is posteriorly wider (Cx-I basal width/Cx-I+II width: robustipalpis male 0.36���0.37, female 0.40; intermedius male 0.34���0.35, female 0.36); the palps (especially P-4) are more compact: P-4 L/H 2.5���2.7 (robustipalpis male), 2.5���2.6 (robustipalpis female); 3.0���3.1 (intermedius male) 3.1 (intermedius female). Hygrobates robustipalpis sp. nov. (especially the male) is the smallest Lurchibates species so far described: idiosoma L/W 640���670/560���600 (robustipalpis male); 760���920/580���720 (other species of Lurchibates males). The female of H. intermedius sp. nov. is similar to H. aloisii, especially in the shape of the genital field, as in both species the genital plates are flanking the posterior 2/3 of the genital opening; however, in the latter species the genital plates are anterior more pointed. In H. salamandrarum the genital plates are in the same position as in the two species mentioned before, but this species is characterized by a very broad gnathosoma and a very short P-4: L/H 2.1 (H. salamandrarum, female); 3.0 (intermedius sp. nov., female); 3.0���3.2 (H. aloisii, female). As mentioned before, H. robustipalpis sp. nov., the smallest species described so far, is characterized by the combination of it���s dumbbell-shaped genital field, relatively compact palps and a dense, rather large denticle field ventrally at P-3 (as well as relatively many denticles ventrally at P-2); the female is separated from those of the other species of the subgenus mainly by it���s typical palp (similar to the male���compact P-4 (L/H 2.5���2.6), large strong claws and large pointed basal cone at P-5, dense field of denticles at P-3) and relatively large, broad genital plates larger than the genital opening. Hygrobates pilosus sp. nov. is mainly characterized by the very characteristic male genital field���it���s wing-like shape and large number of setae is clearly separating the species from all other so far described ones; the anterior coxal group (with the pointed, narrow posterior margin) is slightly similar to H. forcipifer, however it���s rectangular Cx-IV, and especially the shape of the genital field, are clearly separating H. pilosus sp. nov. from the latter. There are also obvious differences in the shape of the genital skeleton, with regard to the poor conditions of this structure in most preparations so far these are not used systematically in the separation of the species., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F. & Shimano, Satoshi, 2020, Description of three new water mite species of Hygrobates Koch, 1837 (Lurchibates Goldschmidt & Fu, 2011) (Acari, Hydrachnidia, Hygrobatidae), parasitic in newts of the genera Paramesotriton and Pachytriton (Amphibia, Caudata, Salamandridae) from China, pp. 25-42 in Zootaxa 4768 (1) on pages 36-41, DOI: 10.11646/zootaxa.4768.1.3, http://zenodo.org/record/3777804

Hygrobates robustipalpis sp. nov. Goldschmidt, Nishikawa & Shimano Material examined: Holotype male, slide mounted in glycerine jelly, preparation no. CIB INV 0004, parasitic on Pachytriton inexpectatus (Amphibia, Caudata, Salamandridae): newt was collected in China, Guizhou Province, Leishan County, Mt. Leigong on September 24th 2011, preserved in 70% Ethanol; mite was attached to the fore- or hindlimb of the newt (CIB GZ 20110948). Paratypes: two females CIB INV 0005 from mouth cavity, limbs, or lateral body of the newt (CIB GZ 20110947) and CIB INV 0006 from mouth cavity, axilla, groin (base of the hindlimb), or lateral body of the newt (CIB GZ 20110948), one male CIB INV 0007 from mouth cavity, limbs, or lateral body of the newt (CIB GZ 20110947)��� newts data are the same as holotype���s host. Distribution: All specimens of H. robustipalpis sp. nov. have been collected from Pachytriton inexpectatus from Guizhou Province, China. The new species is at maximum limited to the same distribution as its host, which is so far known from Eastern Guizhou, southwestern and southern Hunan, extreme northwestern Guangdong, and northern and eastern Guangxi, China. Derivatio nominis: robustipalpis ���derived from robustus (lat.)���strong, powerful; palpus (lat.)���palp; named for the very strong and pointed basal cone and the long and pointed terminal claws at P-5. Diagnosis: Characters of the subgenus; idiosoma relatively small, coxal field relatively broad (compared to most other species of the group), anterior coxal group posteriorly broad; male genital field rounded, apple-shaped, with many setae regularly arranged at outer margin; female genital plates broad kidney-shaped, larger than genital opening, shifted posteriorly; distal leg segments relatively shorter than in other species; P-2 ventro-distally bearing a few denticles, P-3 ventro-distally bearing many denticles, P-4 relatively compact, proximo-ventral extension of P-5 large, pointed, tip slightly curved ventrally; cheliceral claw relatively straight, long and slender. Description, Male (n = 2): Idiosoma rounded-oval, L/W ratio 1.12 (1.13), L/W 672 (636)/600 (564); fused anterior coxae of both sides slender, narrow triangular (Cx-I + II L/W 294 (277)/462 (402)), medio-posterior margin extended by secondary sclerotization (posteriorly convex, laterally curved, pointed); Cx-I basal width 176 (153), gnathosoma broad, posteriorly slightly converging, lateral separation reaching far posteriorly, widely fused with the posterior part of the first coxae, anterior tip of gnathosoma slightly projecting (Fig. 20); posterior coxal groups (Cx- III + Cx-IV of one side) L/W 247 (228)/252 (197), anterior and posterior coxal groups laterally very close, medially diverging (however, less than given in Fig. 20, as in the type-specimen the integument is broken and the posterior coxal groups are slightly shifted to lateral as an artifact); posterior coxae obtuse triangular medial edges rounded, formed by Cx-IV only, Cx-IV short and broad, trapezoid, posterior margin straight transversely, lateral margin straight (Fig. 20); genital field rounded apple-shaped, acetabular plates anteriorly and posteriorly overlapping the pre- and postgenitale, with 24/27 (27/30) setae regularly arranged at outer margin on each side (Fig. 20), genital opening elongate-oval, genital field L/W 192 (144)/ 296 (258); acetabula irregularly elongated oval (L/W Ac-1, 104 (57)/49 (36), Ac-2, 98 (83)/48 (39), Ac-3, 98 (60)/42 (43)); genital skeleton partly broken in type specimen (completely destroyed in second specimen), as far as visible shape typical for the subgenus, brachia distalia very long and slender, regularly curved (Fig. 21); all legs slender, bearing many heavy setae, dorso-distally at I- and II-leg-1 and -2 with groups of four to six heavy setae, distal segments straight (Figs. 22���25); measurements (L/H) of distal leg segments: I-leg-5, 234 (210)/48 (48), I-leg-6, 192 (192)/48 (46); II-leg-5, 234 (228)/48 (48), II-leg-6, 198 (198)/48 (42); III-leg-5, 258 (252)/50 (48), III-leg-6, 222 (216)/54 (48); IV-leg-3, 168 (156)/66 (58), IV-leg-5, 270 (258)/ 54 (48), IV-leg-6, 234 (223)/48 (42); chelicerae very strong, with a relatively short, high basal segment and long basal groove; cheliceral claws large (slightly shorter than in other species), slender, distally straight, sharply pointed, dorsal margin in the distal half with strong serration that continues proximally in a lateral serration, medially and laterally striated (Figs. 26, 27); palps strong (Figs. 28, 29), ventral margin of P-2 slightly concave, antero-ventral corner bearing a few denticles, P-3 ventro-laterally with many large, pointed denticles in distal 2/3, P-4 compact, straight, distally slightly curved, with a pair of ventral setae in the distal third, distal margin strongly rotated ventrally; P-5 ventro-proximally bearing a well developed, pointed, cone-shaped projection, with a ventrally-curved, sharp tip, dorso-distal seta compact, denticle-like, sticking out, ventro-distal pair of large claws long and pointed; mouthpart measurements: chelicera total L 372 (353), maximum H 102 (102), L/H ratio 3.7 (3.5), basal segment L 235 (216), claw L 183 (167), basal segment/claw ratio 1.3 (1.3); curvation of cheliceral claw 22�� (23.5��), palp total L 451 (423), L/H P-1, 52 (42)/73 (68), P-2, 127 (115)/85 (85), P-3, 80 (80)/78 (75), P-4, 146 (139)/59 (52), P-5, 47 (47)/54 (47). Description, Female (n = 2): Idiosoma similar to male, much larger, L/W 816���1380/684���1296, and more rounded (L/W ratio 1.06���1.19); unpaired anterior coxal group as in males, L/W 312���336/462���486, medio- and latero-posterior margin extended by secondary sclerotization, with slightly curved lateral tips; Cx-I basal width 186���193; gnathosoma anteriorly more curved, slightly more projecting than in males, separation to Cx-I posteriorly converging, reaching far posteriorly (as in males) (Fig. 30); paired posterior coxal groups closer to each other and to the anterior coxal group than in males, more slender, posterior margin inclined from postero-lateral to antero- medial, triangular in shape with medial edges formed by Cx-IV only, L/W 270���294/252���259 (Fig. 30); genital field relatively wide and large (broader than in males), with strongly sclerotized pre- and postgenitale, genital plates well distant from each other, posteriorly by far more approached than anteriorly, separated by soft integument, broad kidney-shaped, anteriorly rounded, clearly longer than genital opening, level of anterior margin lightly posterior to pregenitale, posterior margin extending far beyond postgenitale (Fig. 30), total genital field L/W 204���282/318���366; acetabula irregularly-oval (some rather triangular or rectangular) (L/W Ac-1, 79���85/42, Ac-2, 91���98/44���46, Ac- 3, 86���95/45���49); 21���26 setae arranged mainly at the anterior, posterior and lateral margins of the genital plates, one or two pairs anterior to genital plates (Fig. 30); legs similar to males (Figs. 31���34) measurements of distal leg segments: L/H I-leg-5, 238���240/49���50, I-leg-6, 204���216/48; II-leg-5, 252���262/54���55, II-leg-6, 216���222/46���55; III-leg-5, 283���284/54���55, III-leg-6, 235���242/48���54; IV-leg-3, 180���198/62���68, IV-leg-5, 294���300/54���58, IV-leg- 6, 240���264/42���54; chelicerae similar to males, slightly stronger (Figs. 35, 36); palps similar to males (Figs. 37, 38); mouthpart measurements: chelicera total L 420���432, maximum H 120���124, L/H ratio 3.5, basal segment L 268���277, claw L 198���209, basal segment/claw ratio 1.3���1.4; cheliceral claw curve 21���22��; palp total L 479���498, L/H P-1, 52���54/80, P-2, 132���139/87���94, P-3, 94���99/82���85, P-4, 153���155/59���61, P-5, 49���52/59���61., Published as part of Goldschmidt, Tom, Nishikawa, Kanto, Hiruta, Shimpei F. & Shimano, Satoshi, 2020, Description of three new water mite species of Hygrobates Koch, 1837 (Lurchibates Goldschmidt & Fu, 2011) (Acari, Hydrachnidia, Hygrobatidae), parasitic in newts of the genera Paramesotriton and Pachytriton (Amphibia, Caudata, Salamandridae) from China, pp. 25-42 in Zootaxa 4768 (1) on pages 31-35, DOI: 10.11646/zootaxa.4768.1.3, http://zenodo.org/record/3777804

Supplement 1. Story of discovering the “undescribed species” on Twitter (by SS, one of authors) with screenshots of TN’s (one of authors) four tweets on PC browser.

Trichogalumna ekaterinae sp. nov. (Fig. 1) Diagnosis. Small species, body length 253���262 ��m, width of notogaster 173���182 ��m; rostrum widely rounded in dorsal view, but protruding in lateral view; rostral, lamellar and interlamellar setae medium long, thin, smooth; sensillus with thin, long stalk and widely lanceolate head with small, pointed tip; anterior margin of notogaster not developed; notogastral setae well developed; three pairs of porose areas (Aa, A 1 and A 3). Measurements. Holotype (female): body length of 253 ��m, width of notogaster 176 ��m; paratypes (three females): 256���262 (259) ��m, width of notogaster 173���182 (178) ��m. Integument (Figs. 1A). Body color yellowish brown to dark brown. Body surface nearly smooth, with microgranules on prodorsum and pteromorph. Prodorsum (Fig. 1A, B, C, E). Rostrum widely rounded in dorsal view, but protruding in lateral view; inner rostral tooth (irt) of rostrum distinct (Fig. 1B). Tutorial (S) and lamellar (L) ridges distinct, nearly parallel, curving backwards. Rostral (ro) and lamellar (le) setae 30���32 ��m in length, interlamellar seta (in) ��� 22 ��m long, setiform, thin, smooth. Sensillus (ss) having thin stalk and large, asymmetrically dilated, almost semicircular head with small, pointed tip. Dorsosejugal porose area (Ad) oval, located posterolaterad to seta in. Exobothridial seta and porose area Al not evident. Notogaster (Fig. 1A). Anterior margin of notogaster not developed. Dorsophragma (D) elongated longitudinally. Notogastral setae 18���32 ��m long, thin, smooth. Three pairs of notogastral porose areas, Aa, A 1 and A 3 circular to oval in shape (in holotype, left A 1 was divided into two parts); A 1 located anterior to seta h 2; A 3 situated posterior to seta h 1. Notogastral lyrifissures (ia, im, ih, ips and ip) and opisthonotal gland opening (gla) well developed; ip visible in posterior view. Gnathosoma (Fig. 1D). Morphology of subcapitulum, palp and chelicera typical for genus (e.g. see Ermilov & Corpuz-Raros 2016). Subcapitular setae short, setiform, smooth. Palp setation: 0-2-1-3-9+1��; axillary saccules distinct, elongated. Cheliceral setae setiform, barbed, cha longer than chb; Tr��g��rdh���s organ elongate triangular, rounded distally. Epimeral region (Fig. 1D). Only three pairs of epimeral setae (1a, 3b, 4b) distinct, thin, smooth; seta 1a ��� 21 ��m long, 3b and 4b ��� 11 ��m long. Discidium broadly conical; pedotectum I rounded, pedotectum II scale-like, subtriangular in ventral view. Circumpedal carina well developed. Ano-genital region (Fig. 1D). All ano-genital setae medium long (12-18 ��m), thin, smooth; g 1, g 2 and g 3 inserted on anterior margin of genital plate, g 4, g 5 arranged longitudinally, g 6 inserted on posterior margin of genital plate. Adanal setae ad 1 and ad 2 inserted posterior to anal plates; ad 3 inserted in paraanal position; lyrifissure iad situated anterolateral to anal aperture. Postanal porose area (Ap) oval, transversely oriented, visible only in posterior view. Legs. Morphology of leg segments, setae and solenidia typical for Trichogalumna (e.g. Ermilov et al. 2011). Claws smooth dorsally. Famulus short, with slight distal expansion. Formulas of leg setation: I (1-4-3-4-20), II (1- 4-3-4-15), III (1-2-1-3-15), IV (1-2-2-3-12); formula of solenidia: I (1-2-2); II (1-2���2), III (0-1-1), IV (0-1-0). Material examined. Holotype (female) and three paratypes (females): Raelkelat, Island of Babeldaob, Republic of Palau, from moss sample growing on bark of fallen tree in a secondary forest, 7��28���59���N, 134��29���32���E, coll. S. Shimano, 17 December 2017. Type depository. Holotype and all paratypes are deposited in NSMT���the National Museum of Nature and Science, Tsukuba, Japan (Zhang 2018). Etymology. This species is named in honor of our esteemed colleague and friend Dr. Ekaterina A. Sidorchuk, who made a great contribution to the knowledge of mite diversity in Baltic amber inclusions. Remarks. Trichogalumna seminuda Balogh, 1960 known from Angola, Trichogalumna arborea Ohkubo, 1984 from Japan, Trichogalumna vietnamica Mahunka, 1987 from Vietnam, Trichogalumna africana Ermilov, Sidorchuk & Rybalov, 2011 from Ethiopia, and a semicosmopolitan species, Trichogalumna nipponica (Aoki, 1966) (see Balogh 1960; Aoki 1966; Ohkubo 1984; Mahunka 1987; Ermilov et al. 2011) are similar to Trichogalumna ekaterinae sp. nov. in the morphology of asymmetrically dilated sensillus. However, all these known species have clearly developed four pairs of porose areas (vs. A 1 absent in the new species), and a much larger body size (body sizes more than 350 �� 240 ��m vs. 259 �� 178 ��m). Additionally, T. seminuda differs from the present new species by the very long interlamellar seta extending beyond the rostrum, and distinctly barbed rostral setae. The other African species, T. africana is different from the present new species in the relatively slender sensillus with a small notch behind the tip of the sensillar head. The Japanese species, T. arborea differs from the new species in the well-developed transverse bands on the notogaster and ano-genital region, as well as in the finely wrinkled and granulated structure of the humeral region. The Oriental species, T. vietnamica can easily be distinguished from T. ekaterinae sp. nov. by the long, barbed rostral seta. The semicosmopolitan species, T. nipponica has a much slender, distally barbed lanceolate sensillus rather than the largely dilated, smooth sensillus shown in the present new species. It should be noted here that the holotype of Trichogalumna ekaterinae sp. nov. has the left porose area A 1 divided into two parts, which seems to be an abnormal character. Klimov & Ermilov (2017) studied the evolutionary dynamics of gain and loss of the notogastral porose areas of Galumnoidea Jacot, 1925 with respect to the various modifications of their properties, such as shape and position. These authors concluded that patterns of expression of porose areas in abnormal individuals and rare species suggest that this may be a complex, non-Mendelian character, encoded by several genes (i.e. a polygenic trait). They also proposed that the loss of porose areas is not likely to be down-regulated by a third gene. Genus Galumna Heyden, 1826 Type species: Notaspis alatus Hermann, 1804, Published as part of Bayartogtokh, Badamdorj & Shimano, Satoshi, 2019, Contribution to the knowledge of Galumnidae (Acari: Oribatida) in the Oriental region, pp. 368-377 in Zootaxa 4647 (1) on pages 369-371, DOI: 10.11646/zootaxa.4647.1.23, http://zenodo.org/record/3353188, {"references":["Ermilov, S. G. & Corpuz-Raros, L. (2016) New species and records of Galumnidae (Acari, Oribatida) from the Philippines. Zootaxa, 4171 (1), 77 - 100. https: // doi. org / 10.11646 / zootaxa. 4171.1.3","Zhang, Z. - Q. (2018) Repositories for mite and tick specimens: acronyms and their nomenclature. Systematic & Applied Acarology, 23 (12), 2432 - 2446. https: // doi. org / 10.11158 / saa. 23.12.12","Balogh, J. (1960) Oribates (Acari) nouveaux d'Angola et du Congo Belge, 2 eme serie. Companhia de Diamantes de Angola, 51, 15 - 40.","Ohkubo, N. (1984) Several species of Trichogalumna (Acarina, Oribatida) from Japan. Acarologia, 25 (3), 293 - 306.","Mahunka, S. (1987) A survey of the Oribatid (Acari) fauna of Vietnam, I. Annales Historico-Naturales Musei Nationalis Hun- garici, 79, 259 - 279.","Ermilov, S. G., Sidorchuk, E. A. & Rybalov, L. B. (2011) Three new species of oribatid mites (Acari: Oribatida: Galumnoidea) from Ethiopia. International Journal of Acarology, 37 (Supplement 1), 2 - 17. https: // doi. org / 10.1080 / 01647954.2010.528799","Aoki, J. (1966) The large-winged mites of Japan (Acari: Cryptostigmata). Bulletin of the National Science Museum, Tokyo, 9 (3), 257 - 275.","Ermilov, S. G. & Klimov, P. B. (2017) Generic revision of the large-winged mite superfamily Galumnoidea (Acari, Oribatida) of the world. Zootaxa, 4357 (1), 1 - 72. https: // doi. org / 10.11646 / zootaxa. 4357.1.1","Jacot, A. P. (1925) Phylogeny in the Oribatoidea. American Naturalist, 59 (662), 272 - 279. https: // doi. org / 10.1086 / 280038","Heyden, C. (1826) Versuch einer systematischen Eintheilung der Acariden. Isis, Oken, 1 (4), 607 - 613.","Hermann, J. F. (1804) Memoire apterologique. F. G. Levrault, Strassbourg, 144 pp."]}

Waki, Tsukasa, Shimano, Satoshi (2019): Redescription of the Snail Mite Riccardoella reaumuri (Acariformes: Prostigmata: Ereynetidae). Species Diversity 24: 97-102, DOI: 10.12782/specdiv.24.97

Waki, Tsukasa, Shimano, Satoshi, Asami, Takahiro (2019): First Record of Riccardoella (Proriccardoella) triodopsis (Acariformes: Trombidiformes: Ereynetidae) from Japan, with Additional Morphological Information. Species Diversity 24 (1): 11-15, DOI: 10.12782/specdiv.24.11, URL: http://dx.doi.org/10.12782/specdiv.24.11

Genus Riccardoella Berlese, 1923 Type species: Riccardoella limacum (Schrank, 1776), Published as part of Waki, Tsukasa & Shimano, Satoshi, 2018, A new species of the genus Riccardoella (Acari: Prostigmata: Ereynetidae) from the land snail Tauphaedusa tau (Gastropoda: Clausliidae) in Japan, pp. 163-174 in Zootaxa 4402 (1) on page 164, DOI: 10.11646/zootaxa.4402.1.8, http://zenodo.org/record/1208315, {"references":["Berlese, A. (1923) Centuria sesta di Acari nuovi. 1. Prostigmata. Redia, 15, 242 - 246."]}