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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]"]}