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Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. urn:lsid:zoobank.org:act: 40C3DB96-19CD-4354-ACA9-434F0ADA34BB Figs 2���12 Diagnosis Troglomorphic species. Antennae with 39���45 antennomeres; 12 complex olfactory chemoreceptors within cupuliform organ; non-protruding frontal process slightly protruding, plain, with non-tubercular setae or just slightly tubercular. Pronotum 1+1 ma, 2+2 la 1,2 , 2+2 lp 2,3 ; mesonotum with 1+1 ma, 3+3 la 1���3 , 2+2 lp 2,3 , 1+1 mp; metanotum with 1+1 ma, 2+2 la 1,2 or sla 1,2 , 1+1 lp 2 , 1+1 mp; thin; all notal macrosetae long and covered by thin barbs on half to distal portions; thin, middle-sized clothing setae covered by 0���4 thin distal barbs. Legs elongated, pretarsus of metathoracic legs slightly overpasses end of abdomen. Femora I���III with one long, barbed dorsal macroseta and one shorter, barbed ventral macroseta. Tibiae I���III with two short barbed ventral macrosetae. Claws unequal (posterior claw 1.3 �� as long as anterior one); large, backward overhang on posterior claw; lateral crests well-developed. Pretarsal process long and setiform. Urotergites 1+1 post 1 on I���II; 0+0, 0+1 or 1+1 la, 1+1 or 2+2 post 1,2 on III; 1+1 la 3 , 2+2 to 4+4 post 1���4 on IV; 2+2 la 2,3 , 4+4 post 1���4 on V���VII; 6+6 post 1���6 on VIII and 8+8 or 8+7 post 1���8 on abdominal IX. Urosternite I with 8+8���7+7 macrosetae (Figs 8���9); urosternites II���VII with 6+6 macrosetae; urosternite VIII with 2+2 macrosetae. Male urosternite I (Fig. 8) with slightly enlarged subcylindrical appendages, each bearing up to 21 glandular a 1 setae. Female appendages slightly thinner, with up to 11 glandular a 1 setae. Etymology The specific epithet refers to the Imereti region, the location of the Shvilobisa Cave, treated as a noun in apposition. Type material Holotype GEORGIA ��� ♀, ���holotype-♀ IZISU-TD-T-00001���; Shvilobisa Cave, Bunikauri village, Chiatura Municipality, Imereti region, Zemo Imereti Plateau; 42��19���31.44��� N, 43��16���4.33��� E; 24 Feb. 2018; Shalva Barjadze and Eter Maghradze leg.; IZISU-TD-T-00001. Paratypes GEORGIA ��� 1 ♂, ���paratype-♂1 IZISU-TD-T-00002���; same collection data as for holotype; IZISU- TD-T-00002 ��� 1 ♀, ���paratype-♀1 MZB (MCNB) 2021-2336���; same locality as for holotype; 20 Jul. 2020; Eter Maghradze leg.; MZB (MCNB) 2021-2336 ��� 1 ♀, ���paratype-♀2 MZB (MCNB) 2021-2337���; same collection data as for preceding; MZB (MCNB) 2021-2337 ��� 1 ♂, ���paratype-♂2 Coll AS���; same collection data as for preceding; Coll AS. Other material GEORGIA ��� 2 specs, unknown sex [for SEM photography and one specimen for DNA analysis]; Shvilobisa Cave; 24 Feb. 2018; Shalva Barjadze and Eter Maghradze leg. ��� 2 specs [for SEM photography and one for DNA analysis]; same collection data as for preceding; 20 Jul. 2020; Eter Maghradze leg. Other material from two other caves (all Coll AS) GEORGIA ��� 1 ♀; Kumistavi village Tskaltubo Municipality, Imereti Region, Sataplia-Tskaltubo karst massif, Datvis (Bear) Cave; 42��22���28������ N, 42��35���45��� E; 5 Jul. 2018; Eter Maghradze leg. ��� 1 ♀; same collection data as for preceding; 1 Sept. 2019 ��� 1 ♂; near Melouri village, Tskaltubo Municipality, Imereti Region, Sataplia-Tskaltubo karst massif, Melouri Cave; 42��23���15.1��� N, 42��37���41.5��� E; 1 Nov. 2018; Eter Maghradze leg. Description BODY. Body length 4.3���7.2 mm (females) and 4.9���5.2 mm (males) (Table 1). Epicuticle smooth under optical microscope and SEM; body with thin, middle-sized clothing setae covered by 0���4 thin distal barbs. HEAD. Three intact antennae, all slightly longer than body length, with 39���45 antennomeres (Table 1). Small, thin, subcylindrical sensillum on third antennomere located in ventral position between c and d macrosetae. Central antennomeres 2.1 �� as long as wide, apical antennomere 3.0 �� as long as wide. Cupuliform organ occupying ⅓ of total length of apical antennomere, with about 12 complex olfactory chemoreceptors. Each olfactory chemoreceptor is composed of a complete fold surrounding a central cylinder with two lateral expansions, entirely reticulated and perforated (Fig. 5). Gouge sensilla 30��� 40 ��m long, in a single distal whorl of 13���16 sensilla on each medial and distal antennomere. Frontal process slightly protruding, plain, with non-tubercular setae or just slightly tubercular on distal portion (Fig. 3); macrosetae along the insertion line of antennomere and i macrosetae and x setae longer than other macrosetae (a / i / p/ x with relative lengths of 25 / 36 / 19 / 37 in holotype). Suboval labial palps with a bacilliform latero-external sensillum, two guard setae, up to 7 setae on anterior border, and up to 130 neuroglandular setae in holotype. THORAX. Thoracic macrosetal distribution (Figs 2, 4, 6): pronotum with 1+1 ma, 2+2 la 1���2 , 2+2 lp 2,3 ; mesonotum with 1+1 ma, 3+3 la 1���3 , 2+2 lp 2,3 , 1+1 mp; metanotum with 1+1 ma, 2+2 la 1,2 or sla 1,2 , 1+1 lp 2 , 1+1 mp. All notal macrosetae are long and covered by thin barbs on half to distal portions (Figs 4, 6); submacrosetae sla are thinner and shorter than notal macrosetae; marginal setae are similar to clothing setae, and covered by 1���8 thin distal barbs. Legs elongated, pretarsus of metathoracic legs slightly overpasses end of abdomen (Table 1). Femora I���III with one long, barbed dorsal macroseta and one shorter, barbed ventral macroseta. Tibiae I���III with two short barbed ventral macrosetae. Calcars with 4���5 long barbs. Tarsi with two rows of thicker ventral setae with 2���3 very thin barbs on middle portion. Two dorsal and one ventral, smooth, subapical tarsal setae. Claws are unequal (posterior claw 1.3 �� as long as anterior one); large, backward overhang on posterior claw; lateral crests well-developed. Pretarsal process long and setiform, overpassing end of claws. ABDOMEN. Distribution of abdominal macrosetae on tergites (Fig. 7): 1+1 post 1 on I���II; 0+0, 0+1 or 1+1 la, 1+1 or 2+2 post 1,2 on III; 1+1 la 3 , 2+2 to 4+4 post 1���4 on IV; 2+2 la 2,3 , 4+4 post 1���4 on V���VII; 6+6 post 1���6 on VIII and 8+8 or 8+7 post 1���8 on abdominal segment IX. All post urotergal macrosetae long and covered by thin barbs along distal four-fifths (Fig. 10); la urotergal macrosetae shorter than post macrosetae, covered by barbs along distal half. Urosternite I with 8+8���7+7 macrosetae (Figs 8���9); urosternites II���VII with 6+6 macrosetae; urosternite VIII with 2+2 macrosetae (Fig. 12); all urosternal macrosetae robust and large, covered by long barbs along distal third to four-fifths. Apical, subapical and ventromedial setae with a few (two to four) thin, short and long barbs (Fig. 11). SECONDARY SEX CHARACTERS. Male urosternite I (Fig. 8) with slightly enlarged subcylindrical appendages, each bearing up to 21 glandular a1 setae. Female appendages slightly thinner, with up to 11 glandular a 1 setae. Spermatozoid fascicles 40 ��m in diameter without apparently spiral filament. Molecular analysis The nucleotide substitution model selected was GTR+G+I (BIC = 6998.6), with the proportion of invariant sites (I = 0.46) and estimated alpha parameter for the gamma distribution (�� = 1.39), indicating a significant heterogeneity in the DNA substitution among sites. The Campodeidae sequences formed a well-supported clade, clearly distinct from that of Japygidae (Fig. 13). Although bootstrap values were low, the ML phylogenetic tree grouped Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. with Eastern Europe taxa such as Plusiocampa (Plusiocampa) aff. elongata Ionescu, 1955 and Plusiocampa (Plusiocampa) humicola Ionescu, 1955, whereas Iberian Peninsula taxa (Plusiocampa (Plusiocampa) gadorensis Sendra, 2001, Plusiocampa (Plusiocampa) baetica Sendra, 2004 and Cestocampa iberica Sendra & Cond��, 2012) clustered in a distinct clade. K2P genetic distances also showed P. (P.) aff. elongata (0.206�� 0.027) and P. (P.) humicola (0.205 �� 0.028) to be the closest species to the new Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. from Georgia. Habitat Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. inhabits the deep zone (over 50 m from the entrance) of three caves. The Shvilobisa Cave, the type locality, is a 1000 m long, tunnellike, easily accessible sub-horizontal cave with a small subterranean water stream (Tatashidze et al. 2009b). The others two nearby caves are about 55 km away from the Shvilobisa Cave; the Melouri Cave is 5300 meters long and has the status of natural monument (Tatashidze et al. 2009b), whereas the Datvis Cave is a poorly known cavern (K. Tsikarishvili, pers. comm.). The distance between the Datvis and Melouri caves is ca 3.5 km. The Melouri Cave ��� easily accessible ��� has dried halls and a permanent subterranean water stream near its end. This cave has gigantic stalagmites and fallen stones. The Datvis Cave is a horizontal, dry, and easily accessible cave with several halls, which are rich in different speleothems like the Shvilobisa Cave (Tatashidze et al. 2009b). Invertebrate cavernicolous species of the studied caves The three caves (Shvilobisa, Datvis, and Melouri) which Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. inhabits are also the dwellings of other troglobitic arthropod species. Datvis and Melouri caves share three troglobitic species: the Diplopoda Leucogeorgia prometheus Antić & Reip, 2020, the Isopoda Colchidoniscus kutaissianus Borutzky, 1974, and the Pseudoscorpionida Chthonius satapliaensis Schawaller & Dashdamirov, 1988. In addition, the Insecta (Carabidae Coleoptera) Troglocimmerites imereti Dolzhanski & Ljovuschkin, 1985 dwells in the Datvis cave; and the Melouri Cave has five more species: the Opiliones Nemaspela melouri Martens, Maghradze & Barjadze, 2021, the Araneae Centromerus bulgarianus Drensky, 1931, the Hexapoda Collembola Pseudacherontides zenkevitchi Djanashvili, 1971; the Insecta (Carabidae Coleoptera) Inotrechus kurnakovi Dolzhanski & Ljovuschkin, 1989; and Troglocimmerites sp. 1. In the Shvilobisa Cave, eight troglobitic species dwell: the Diplopoda Leucogeorgia gioi Antić & Reip, 2020, the Isopoda Caucasonethes cf. borutzkyi Verhoeff, 1932 and Colchidoniscus sp., the Pseudoscorpionida Chthonius satapliaensis Schawaller & Dashdamirov, 1988, the Opiliones Nemaspela sp., the Hexapoda Collembola Oncopodura sp. and Pseudosinella sp., and the Insecta (Carabidae Coleoptera) Troglocimmerites sp. 2. (Barjadze et al. 2019; Maghradze et al. 2019; Antić & Reip 2020; Martens et al. 2021; Maghradze & Barjadze, unpublished data)., Published as part of Sendra, Alberto, Palero, Ferran, S��nchez-Garc��a, Alba, Jim��nez-Valverde, Alberto, Selfa, Jes��s, Maghradze, Eter & Barjadze, Shalva, 2021, A new Diplura species from Georgia caves, Plusiocampa (Plusiocampa) imereti (Diplura, Campodeidae), with morphological and molecular data, pp. 71-85 in European Journal of Taxonomy 778 on pages 74-79, DOI: 10.5852/ejt.2021.778.1567, http://zenodo.org/record/5675209, {"references":["Ionescu M. A. 1955. Diplura. In: Academia Republicii Socialiste Romania (ed.) Fauna Republicii Populare Romane, Insecta 7 (2): 1 - 48.","Sendra A., Lara M. D., Ruiz Aviles F. & Tinaut A. 2004. Une nouvelle espece du genre Plusiocampa Silvestri, 1912 (Diplura, Campodeidae) et donnees pour sa reconstruction paleobiogeographique dans les Betiques. Subterranean Biology 2: 113 - 122.","Tatashidze Z. K., Tsikarishvili K. D. & Jishkariani J. M. 2009 b. The Cadastre of the Karst Caves of Georgia. Petiti Publishing House, Tbilisi [in Georgian].","Antic D. Z & Reip H. R. 2020. The millipede genus Leucogeorgia Verhoeff, 1930 in the Caucasus, with descriptions of eleven new species, erection of a new monotypic genus and notes on the tribe Leucogeorgiini (Diplopoda: Julida: Julidae). European Journal of Taxonomy 713: 1 - 106. https: // doi. org / 10.5852 / ejt. 2020.713","Martens J., Maghradze E. & Barjadze Sh. 2021. Two new species of the genus Nemaspela Silhavy from caves in Georgia (Opiliones: Nemastomatidae). Zootaxa 4951 (3): 541 - 558. https: // doi. org / 10.11646 / zootaxa. 4951.3.7","Barjadze Sh., Arabuli T., Mumladze L., Maghradze E., Asanidze Z. & Shavadze R. 2019. Cave Biodiversity of Georgia, Open Access Database. Institute of Zoology at Ilia State University. Available from https: // cbg. iliauni. edu. ge / en / [accessed: 20 Apr. 2021].","Maghradze E., Faille A., Barjadze Sh. & Hlavac P. 2019. A new cavernicolous species of the genus Bergrothia Reitter, 1884 (Coleoptera, Staphylinidae, Pselaphinae) from Georgia. Zootaxa 4608 (2): 371 - 379. https: // doi. org / 10.11646 / zootaxa. 4608.2.11"]}

Genus Pacificampa Chevrizov, 1978 Diagnosis (based on Chevrizov 1978) Thoracic macrosetae with no more than 3+3 macrosetae on pronotum, 4+4 on mesonotum (2+2 lateral posterior included), and 2+2 on metanotum (1+1 lateral posterior included). One dorsal macroseta on metathoracic femora and one or two on tibia. Subequal elbowed claws with ridges on dorsal side that look like very small lateral crests under optical microscope; without lateral processes. Urotergites V– VII with no more than 1+1 medial posterior, 1+1 lateral anterior and 2+2 lateral posterior macrosetae and on urotergite VIII with no more than 1+1 medial posterior and 3+3 lateral macrosetae. Up to 7+7 macrosetae on first urosternite, 5+5 macrosetae on second to seventh urosternites, and 1+1 macrosetae on eighth urosternite. First urosternite in males with thick appendages with large field of glandular a1 setae., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 22, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Pacificampa wudonghuii Sendra sp. nov. urn:lsid:zoobank.org:act: 447EE93A-F466-4C66-827A-0751A259070F Figs 51–64 Etymology This species is dedicated to Professor Wu Donghui, from the Northeast Institute of Geography and Agroecology (NEIGAE), Chinese Academy of Sciences, Changchun (China), for his enthusiastic contribution and support to the knowledge of Northern China soil and cave biodiversity. Type material Holotype CHINA • ♀; Liaoning Province, Benxi, Huanren, Xianren Dong; 17 May 2019, L. Deharveng, A. Bedos and Wu Donghui leg.; labelled “CHILN19-005-holotype”; NEIGA-diplura-01. Paratypes CHINA • 1 ♀, 1 juvenile, same collection data as for holotype; labelled “CHILN19-005-female paratype”; MZB (MCNB) 2020-1160 • 1 juvenile; same collection data as for preceding; labelled “CHILN19- 005-juvenile paratype”; MZB (MCNB) 2020-1161 • 1 ♀; Liaoning Province, Benxi, Huanren, Pylon cave; 17 May 2019; L. Deharveng, A. Bedos and Wu Donghui leg.; labelled “CHILN19-007-female paratype”; MZB (MCNB) 2020-1164. Other material examined CHINA • 1 ♂, same collection data as for holotype; mounted on an aluminium stage and coated with palladium-gold; Coll. AS. Description BODY. Length 6.5–6.9 mm in females; 3.9 mm in one juvenile. Epicuticle smooth under optical microscope but well reticulated under high magnifications as one can see irregular polygonal structures of variable sizes with scattered external glands either visible or covered with secretion (Figs 51, 53); body with smooth clothing setae. HEAD. Antennae broken in all specimens; central antennomeres with two whorls of distal barbed macrosetae and uneven short setae; in addition, with single distal whorl of up to 8–12 gouge sensilla of 22–29 µm long (Fig. 51) and among them one or two small coniform sensilla (5 µm long). Proximal antennomeres with typical trichobothria disposition with bacilliform sensillum (9–10 µm long) on third antennomere in ventral position, between c–d macrosetae (Fig. 52). Plain frontal process with one anterior macrosetae, longer than clothing setae. Three macrosetae along each side of insertion line of antennomere and x setae with length ratios of a / i / p / x, 42/55/42/45, respectively, in holotype (Fig. 54). Large suboval labial palps, each with enlarged coniform latero-external sensillum near two gard setae and eight normal setae on anterior portion, up to 150 neuroglandular setae in medial and posterior positions. THORAX. Thoracic macrosetae distribution: pronotum has 1+1 ma, 1+1 la, 1+1 lp macrosetae; mesonotum has 1+1 ma, 1+1 la and 2+2 lp macrosetae; and metanotum has 1+1 ma and 1+1 lp macrosetae. Long macrosetae with long barbs in distal three-fourths; marginal setae longer than clothing setae, which are barbed from distal half to three-fourths. Legs elongated, metathoracic legs reach posterior border of seventh abdominal segment. Mesothoracic and metathoracic femora have one dorsal macroseta each, barbed along distal three-fourths (Fig. 55), absent in prothoracic femora. Calcars with two or three long barbs in middle (Fig. 56). Prothoracic and mesothoracic tibia with one short ventral macrosetae with one apical barb and two in metathoracic tibia (Fig. 57). Each tarsus with two separated ventral rows of thicker and longer setae among clothing setae, and a few setiform sensilla (Fig. 58). Three long smooth dorsal tarsal and one ventral setae. Subequal elbowed claws with smooth ventral surface ridged on dorsal side that can be mistaken for lateral crests under optical microscopes, between a blunt unguiculus and without lateral processes (Figs 59–60). ABDOMEN. Distribution of abdominal macrosetae on tergites (Fig. 61 shows 1+1 ma on III in the paratype from Grotte du Pylone, but absent in Xianren Dong Cave types; 1+1 ma, 1+1 la and 2+2 lp on IV–VII; 1+1 mp and 3+3 lp on VIII and 1+1 mp and 5+5 lp on IX abdominal; ma and la macrosetae with barbs in distal half to one third and shorter than mp and lp macrosetae, which bear long barbs along distal four-fifths. Urosternite I with 7+7 macrosetae (Fig. 62); urosternites II to VII with 4+4 macrosetae (Fig. 63); urosternite VIII with 1+1 macrosetae (Fig. 64); urosternal macrosetae of varying lengths, with one apical to twenty distal long barbs. Stylus setae smooth, apical and subapical setae shorter than ventromedial seta (Fig. 63). SECONDARY SEX CHARACTERS. Female urosternite I with subcylindrical appendages, each bearing up to 40 glandular a 1 setae in apical field. Male urosternite I with thick short, and subcylindrical appendages, each with large apical field of about 220 glandular a 1 setae (Fig. 62). Taxonomic affinities Pacificampa wudonghuii sp. nov. shares with P. birsteini Chevrizov, 1978, P. caesa Chevrizov, 1978 and P. nipponica Sendra, 2018 the same distribution of notal macrosetae (1+1 medial anterior, 1+1 lateral anterior, and 1+1 lateral posterior macrosetae on pronotum; 1+1 medial anterior, 1+1 lateral anterior, and 2+2 lateral posterior macrosetae on mesonotum; and 1+1 medial anterior and 1+1 lateral anterior macrosetae on metanotum). An undescribed species mentioned by Ferguson (1997) from northeastern China also has this setal distribution. However, P. wudonghuii sp. nov. differs from other species of Pacificampa in having 1+1 medial anterior, 1+1 lateral anterior, and 2+2 lateral posterior macrosetae on the fourth urotergite (Fig. 61), a pattern distribution that begins on the fifth urotergite for the other species. In addition, the larger male appendages have more abundant glandular a 1 setae in P. wudonghuii sp. nov. (Fig. 62) than in the other species of Pacificampa.

Anisuracampa ywangana Sendra & Komerički sp. nov. urn:lsid:zoobank.org:act: 511F81B5-6CB2-4A98-AB3E-11AF72205A26 Figs 1, 3–19; Tables 1–2; Supp. file 2 Etymology This species is named after the important karst region Ywangan in Myanmar, the location of Win Twin Cave (Figs 1, 3–4). Type material Holotype MYANMAR • ♀; Shan State, Ywangan, Win Twin Cave; 21º11′51.47″ N, 96º32′40.34″ E; 16 Nov. 2018; Aung Lin leg.; labelled “holotype”; MZB (MCNB) 2020-1152. Paratypes MYANMAR • 1 ♀; same collection data as for holotype; labelled “paratype-H1”; MZB (MCNB) 2020- 0620 • 1 ♀, 1 ♂, 1 juvenile; same collection data as for holotype; 8 May 2018; labelled “paratype-H2”, “paratype-M”, “paratype-J”; MZB (MCNB) 2020-1153, MZB (MCNB) 2020-1155, MZB (MCNB) 2020-1154. Other material examined MYANMAR • 2 specs; same collection data as for the holotype; 8 May 2018; mounted on two separate aluminium stages and coated with palladium-gold; Coll. AS. Description BODY. Length is 4.3–6.0 mm (n = 4) in adults and 2.6 mm in one juvenile (Table 1). The epicuticle is smooth under optical microscope but reticulated at higher magnification, showing irregular polygonal structures of variable sizes (Fig. 11). Body with long smooth clothing setae. HEAD. Two complete and intact antennae in holotype and paratype-M with 35 antennomeres each, whereas paratype-J (juvenile) has 38 antennomeres. The antennae are similar to length of body (Table 1), with medial antennomeres 2× as long as wide but apical antennomeres 1½ × as long as wide. Cupuliform organ with up to 17 spheroidal olfactory chemoreceptors arranged in two uneven concentric circles; each chemoreceptor forms a complex structure of multi–perforated folds with one distinguishing crown of fringes a surrounding central column. Each of these structures is inside a polygonal cell (Figs 5–6). Distal and central antennomeres with three whorls of distal barbed macrosetae and 5–6 scattered whorls of smooth setae in addition to single distal whorl of up to 14 gouge sensilla of 24–28 µm length (Figs 7–8) that are more abundant on dorsal side of antennomere. Proximal antennomeres with typical trichobothria, plus small bacilliform sensillum on third antennomere in ventral position. Round protrusion of frontal process covered with one anterior macroseta and two or three posterior macrosetae (Fig. 10). Three macrosetae along each side of insertion line of antennomere and x setae with thin distal barbs; length ratios of a / i / p / x, 45/57/56/65, respectively, in paratype-H2. Labial palps large and suboval, each with bacilliform latero-external sensillum, two guard setae, up to 18 normal setae and up to 140 neuroglandular setae. THORAX. Thoracic macrosetae distribution (Figs 9, 11) have pronotum with 1+1 ma, 1+1 la, 2+2 lp 2,3 macrosetae; mesonotum with 1+1 ma, 2+2 la and 4+4 lp 1–4 macrosetae; and metanotum with 1+1 ma, 2+2 la and 3+3 lp 1–3 macrosetae. All macrosetae long and with thin barbs along basal half to twothirds of each seta; marginal setae barbed and longer than clothing setae. Legs elongated, metathoracic legs reach end of abdomen in adults or overpass it in juvenile. Tibia always longer than femur or tarsus (Table 1). Each femur III with three or, less frequently, two long dorsal macrosetae with distal barbs (0.25 mm in 0.62 mm femur of paratype-H1) (femora I–II with one to three long dorsal macrosetae). Calcars covered with thin abundant barbs all over. Tibiae I–III usually with two short, completely barbed ventral macrosetae, occasionally just one. Each tibia with two rows of ventral barbed setae almost from base and scattered throughout with thin, long setiform sensilla. Three well-barbed dorsal distal tarsal setae longer than rest of tarsal setae. Subequal claws with lateral expansion in crest; basal and ventral portion of claws covered with short to long spiniform formations. Laminar pretarsus of lateral processes sharply curved with what apparently looks like thin fringes under optical microscope, but under SEM seen as narrow laminar expansions (Figs 12–15). ABDOMEN. Distribution of macrosetae on tergites with 1+1 post 1 on I–III; 1+1 post 1 to 3+3 post 1–3 on IV, 4+4 post 1–4 and 1+1 la on V, 5+5 post 1–5 or 6+6 post 1–6 and 1+1 la on VI–VII; 7+7 post on VIII; and 8+8 or 9+9 post macrosetae on IX abdominal segment. All tergal abdominal macrosetae long and barbed along distal half to four-fifths. Urosternite I has 7+7 macrosetae (Figs 16–17); while urosternites II to VII with 5+5 macrosetae; and urosternite VIII with 1+1 macrosetae; urosternal macrosetae of medium size and barbed along distal half. Stylus with short apical seta with small barbs, and each subapical seta completely covered with barbs and with short, barbed ventromedial seta (Figs 18–19). One intact cercus 2.4 × body length and with 14 primary articles (Table 2). Article length increases from proximal to distal articles; covered with long thin macrosetae, with distal barbs, and less abundant long, thin setae. Each primary article with whorl of shorter thin plumose setae at distal position. SECONDARY SEX CHARACTERS. Female urosternite I with enlarged subcylindrical appendages, each bearing up to 60 ventral glandular a 2 setae proximally and apical field of up to 28 glandular a 1 setae (Figs 16–17). Male with similar appendages with two fields of glandular setae (19 a 2 and 27 a 1) in one appendage of male paratype. Spermatozoid fascicles small, 0.04 mm in diameter, and formed by undistinguishable filament of spermatozoids in spiral. Taxonomic affinities Anisuracampa was proposed for a soil-dwelling species, Anisuracampa suoxiensis Xie & Yang, 1990 from Hunan Province in southeastern China. As a plusiocampine genus, Anisuracampa is characterized by the laminar pretarsus but with the lateral processes with long barbs and weakly developed lateral crests (Figs 12–15) (see Xie & Yang 1991: fig. 34). In addition to this trait, Anisuracampa has two or three dorsal femoral macrosetae and 1+1 macrosetae on the eighth urosternite. All of these characters are present in A. ywangana sp. nov., a cave-adapted species from the Win Twin Cave in eastern Myanmar. Therefore, these morphological similarities together with the congruent geographical distribution in Southeast Asia are considered sufficient reason to place this new species in Anisuracampa. However, A. ywangana sp. nov. differs from A. suoxiensis in several characters besides the traits related to adaptations to cave ecosystems. Anisuracampa ywangana sp. nov. has 4+4 lateral posterior macrosetae on the mesonotum (Fig. 9) (2+ 2 in A. suoxiensis), 3+3 lateral posterior and 2+2 lateral anterior macrosetae on the metanotum (2+2 lateral posterior and no lateral anterior macrosetae in A. suoxiensis) and 5+5 posterior and 1+1 lateral anterior macrosetae on urotergites V–VII (4+4 posterior and no lateral anterior macrosetae in A. suoxiensis). Regarding the new species’ cave-adapted features, it has a larger body and longer appendages at least double the size of those of A. suoxiensis, with 30–35 antennomeres in A. ywangana sp. nov. (24 in A. suoxiensis). It is impossible to compare other useful taxonomical characteristics mentioned in the description of A. ywangana sp. nov. with A. suoxiensis as the former was described using SEM, whereas the latter was described under optical observations with a brief diagnosis and the type material of A. suoxiensis is not available for study. Remarks Anisuracampa ywangana sp. nov. was observed and collected walking on the Win Twin cave floor and boulders, approximately 300 m from the entrance, and on the wet flowstone in a vast chamber (Figs 1–3; Supp. file 2). The Win Twin cave is located within a large karst area referred to as Ywangan karst, which is situated in the western part of the Shan plateau, approximately 10 km northeast of Ywangan Township and 15 km east of the Panlaung Pyadalin Wildlife Sanctuary. The cave has only been discovered recently and is currently under tourist development by the local community. It has not been fully surveyed and approximately only the first 400 m were explored during the collection of the type material. The cave itself is large, with numerous chambers, and its entrance was enlarged by mining (Fig. 4); after approximately 300 m into the main passage, it opens into a vast chamber filled with speleothems (Fig. 3), from which it again continues further by smaller, narrow passages opening into new chambers. At the bottom of the vast chamber, the oxygen level becomes too low for further exploration during the dry season (May), while by the end of the rainy season (November) it was possible to reach two further lower chambers before the oxygen level became too low. No active water flow was observed, but sparse remains of particular organic matter are present and the air temperature recorded in May 2018 was 20.0°C, while the sediment temperature was 19.6°C (unpublished data)., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on pages 4-9, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Xie R. & Yang Y. 1991. Description of two new genera and three new species of Campodeidae in China (Diplura). Contribution Shanghai Institute of Entomology 10: 95 ‾ 102."]}

Hubeicampa aff. melissa Sendra & Lips gen. et sp. nov. Material examined CHINA • 3 ♂♂ (6.6 to 8.0 mm), 1 ♀ (9.6 mm), 1 juv. (5.5 mm); Hubei, Banqiao, Kedu Dong (Grotte des Têtards); 6 Aug. 2009; Josiane Lips leg.; Coll. AS. Remark Specimens from Kedu Dong are considered as H. aff. melissa gen. et sp. nov. due to the specimens’ bad condition which does not allow us to clearly observe any variations in the taxonomical features. Nevertheless, they have the same important traits as the new species, but there is no dorsal macrosetae on the femur, except in a juvenile. Taxonomic affinities Hubeicampa melissa Sendra & Lips gen. et sp. nov. is closely related to another cave-adapted species, Hubeicampa lipsae (Condé, 1993), which is a new combination of Plusiocampa (Dydimocampa) lipsae Condé, 1993, and thus leads us to propose Hubeicampa as a new genus for both species that live in karst regions of the Chinese province of Hubei. The new genus is distinguished by the pretarsus, which has lateral crests and a laminar lateral process covered by spiniform formations and micro-barbs, pubescent setae and macrosetae (Figs 36–39), as well as by 1+1 macrosetae on the eighth urosternite. A significant number of taxonomic characters separates H. melissa gen. et sp. nov. from H. lipsae. The most relevant differences are in the number of macrosetae on the mesonotum (Fig. 30) (with 1+1 medial anterior, 1+1 lateral anterior and 1+1 medial posterior macrosetae in H. melissa gen. et sp. nov. instead of 1+1 medial anterior and 2+2 lateral posterior macrosetae in H. lipsae), and the number of macrosetae on metanotum (with 1+1 medial anterior macrosetae and none or 1+1 medial posterior submacrosetae instead of 1+1 medial anterior and 1+1 lateral posterior in H. lipsae). Other important features are the larger lateral crests in the pretarsus of H. melissa gen. et sp. nov. (Figs 36–37) the presence of one dorsal macroseta on the metathoracic femora, and one ventral macroseta on the metathoracic tibia in H. melissa gen. et sp. nov. instead of two dorsal femora and two ventral tibia macrosetae in H. lipsae. Finally, there are 4+4 posterior macrosetae on V–VII urotergites in H. melissa gen. et sp. nov. (Fig. 40) instead of 3+ 3 in H. lipsae. Remarks Hubeicampa melissa Sendra & Lips gen. et sp. nov. lives in the “Grotte du 8 ième Ciel Baxian Dong”, a cave situated at an elevation of 1870 m in the mountain near Banqiao village in the Hubei Province. The cave is 1703 m long and 122 m deep, with a huge entrance and giant galleries, as well as narrow passages and a narrow independent entry. The holotype of Hubeicampa melissa gen. et sp. nov. was observed and collected on the cave floor at about 300 m from the huge entry and at the top of the - 40 m deep pit on clay ground above the Saltpetre passage (Lips et al. 2009) (Figs 46–48). Some paratypes were collected in the same cave in the big chamber “Grande galerie du Va-et-Vient”., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on pages 14-18, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Conde B. 1993. Premiers Campodeidae cavernicoles de China, comme exemple de l'evolution souterraine de la Famille (Diplura). Revue suisse de Zoologie 100 (4): 823 ‾ 828. Available from https: // www. biodiversitylibrary. org / page / 41223963 [accessed 19 Nov. 2020].","Lips B., Faverjon M., Routhieau V., Schalk P. & Lips J. 2009. Rapport de la 8 ieme expedition speleologique en Chine, provinces de l'Hubei et du Sichuan. Aventures karstiques lointaines & Institut de technologie de Chengdu. Speleologie au pays de l'Homme sauvage 8: 1 ‾ 112."]}

Genus Hubeicampa Sendra & Lips gen. nov. urn:lsid:zoobank.org:act: 3992F636-8036-4734-ACA6-A4972A157D82 Type species Hubeicampa lipsae (Condé, 1993) comb. nov. Plusiocampa (Dydimocampa) lipsae Condé, 1993: 826, figs 2a–c. Diagnosis Dense pubescence of thin micro-barbs on setae. No more than 4+4 macrosetae on pronotum, 3+3 on mesonotum and 2+2 on metanotum. One or two dorsal macrosetae on metathoracic femora and one or two on tibia. Subequal elbowed claws with lateral crests; basal and ventral portion of claws covered with very small, thin, spiniform formations that look pubescent under optical microscope. Laminar lateral processes of the pretarsus completely covered with dense micro–barbs. Urotergites V–VII with 3+3 or 4+4 posterior macrosetae and without lateral anterior macrosetae. Up to 12+12 macrosetae on first urosternite, 4+4 or 5+5 macrosetae on second to seventh urosternites and 1+1 macrosetae on eighth urosternite. First urosternite in males without glandular g 1 setae and appendages with glandular a 1 setae. Etymology The genus name refers to the Chinese province of Hubei where the first cave-adapted dipluran was described., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on pages 10-11, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Conde B. 1993. Premiers Campodeidae cavernicoles de China, comme exemple de l'evolution souterraine de la Famille (Diplura). Revue suisse de Zoologie 100 (4): 823 ‾ 828. Available from https: // www. biodiversitylibrary. org / page / 41223963 [accessed 19 Nov. 2020]."]}

Genus Hubeicampa Sendra & Lips gen. nov. urn:lsid:zoobank.org:act: 3992F636-8036-4734-ACA6-A4972A157D82 Type species Hubeicampa lipsae (Condé, 1993) comb. nov. Plusiocampa (Dydimocampa) lipsae Condé, 1993: 826, figs 2a–c. Diagnosis Dense pubescence of thin micro-barbs on setae. No more than 4+4 macrosetae on pronotum, 3+3 on mesonotum and 2+2 on metanotum. One or two dorsal macrosetae on metathoracic femora and one or two on tibia. Subequal elbowed claws with lateral crests; basal and ventral portion of claws covered with very small, thin, spiniform formations that look pubescent under optical microscope. Laminar lateral processes of the pretarsus completely covered with dense micro–barbs. Urotergites V–VII with 3+3 or 4+4 posterior macrosetae and without lateral anterior macrosetae. Up to 12+12 macrosetae on first urosternite, 4+4 or 5+5 macrosetae on second to seventh urosternites and 1+1 macrosetae on eighth urosternite. First urosternite in males without glandular g 1 setae and appendages with glandular a 1 setae. Etymology The genus name refers to the Chinese province of Hubei where the first cave-adapted dipluran was described.

Plutocampa speophila Chevrizov, 1978 Plutocampa speophila Chevrizov, 1978: 201, fig. 2(1‾9). Material examined RUSSIA • ♀, holotype (6 mm), 1 ♀ paratype (5 mm); Far East, Primorye region, Olgisk District, Priiskovaya cave; 3 Nov. 1966; S.I. Ljovuschkin leg.; slide; ZMMU Dip0007. Redescription (based on Chevrizov 1978) Under optical microscope it has a smooth cuticle covered by thin smooth clothing setae. Antennae are apparently intact with 23 antennomeres in the paratype, medial antennomeres 1.4 × as long as wide; third antennomere with a thick sensillum in ventral position (c–d macrosetae); last antennomere with at least eight olfactory chemoreceptors. Small and pointed frontal process has one long anterior smooth macrosetae and two shorter posterior macrosetae; there are three smooth macrosetae along each side of the line of insertion of antennomere and smooth x setae with a similar length, ratios of a / i / p / x are 20/23/22/23, respectively. Labial palps are large with thick sensilla. Pronotum with 1+1 ma, 3+3 (2+ 2 paratype) la, 2+2 lp 1,3 mesonotum with 1+1 ma, 3+3 (2+ 2 paratype) la, 2+2 lp 2,3 , 1+1 (0+ 1 paratype) mp and metanotum with 1+1 ma, 2+2 lp, 1+1 mp macrosetae, long and thin notal macrosetae covered with thin barbs along their distal three-fourths; marginal setae are long and some of them are as long and barbed as the macrosetae. The metathoracic legs reach the posterior border of the VII abdominal segment; femora have two long barbed dorsal macrosetae and one ventral macrosetae. The tibia has one ventral short macrosetae, a few distal barbs, and calcars with several long barbs. The tarsus has two rows of smooth ventral setae, slightly thicker than clothing setae and on distal part: three long smooth dorsal setae and one ventral seta; the pretarsus has no lateral processes, with unequal elbowed claws, and with large lateral crests. Urotergite I with 1+1 post 1 ; urotergite II 1+2 (1+ 1 paratype) post 1,2 ; urotergite III 2+2 post 1,2 ; urotergites IV–VII 4+4 post 1–4 ; urotergite VIII 5+5 post 1–5 and abdominal segment IX 7+7 post, macrosetae long with barbs along three-fourths. Urosternite I with 5+5, urosternites II–VII 4+4, and urosternite VIII 1+1 macrosetae; urosternal macrosetae are shorter than urotergal macrosetae with long barbs; smooth stylar setae. First urosternite appendages (female) have large, short and subconical appendages, each with apical field with about fifty glandular a 1 setae., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 20, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Pacificampa caesa Chevrizov, 1978 Pacificampa caesa Chevrizov, 1978: 199, figs 1(10‾19). Material examined RUSSIA • ♂ holotype (6.8 mm); Far East, Primoriye region, Chandolaz Mountain Range, Bliznets Cave; 23 Jun. 1974; B.P. Chevrizov leg.; slide; ZMMU Dip0004. Redescription (based on Chevrizov 1978) It has a smooth cuticle under optical microscope and is covered by abundant short and smooth clothing setae. The antennae are apparently intact with 27 antennomeres, medial antennomeres are 3 × as long as wide; third antennomere has a sensillum in ventral position (c–d macrosetae). The plain frontal process has one long anterior smooth macroseta and two, almost two times shorter, posterior macrosetae; the three macrosetae with short distal barbs along each side of the insertion line of the antennomere are longer than the smooth x setae with length ratios of a / i / p / x, 7/8/4/3, respectively. Pronotum with 1+1 ma, 1+1 la, 1+1 lp, mesonotum with 1+1 ma, 1+1 la, 2+2 lp, and metanotum with 1+1 ma, 1+1 lp macrosetae; all notal macrosetae are robust with a few short distal barbs, in particular the la are longer and the lp pronotal macrosetae have more numerous short distal barbs; the marginal setae are thicker than the clothing setae with short distal barbs, the lateral posterior marginal setae are larger and have more numerous small barbs. The metathoracic legs reach the end of the abdomen; the femora have one robust dorsal macroseta with short distal barbs, whereas the tibia has one ventral short macroseta with 1–2 apical barbs and the calcars are with 3–4 long barbs; the tarsus has two rows of robust, smooth ventral setae, three long, smooth dorsal and one distal ventral setae; the pretarsus is without lateral processes, with subequal elbowed claws, and with a smooth ventral surface ridged on the dorsal side that looks like very small lateral crests under optical microscopes, but they are not. Urotergites V–VII have 1+1 ma, 1+1 la, 2+2 lp; urotergite VIII and abdominal segment IX has 1+1 mp, 3+3 lp, all macrosetae are robust and with a few short distal barbs. Urosternite I has 6+6, urosternites II–VII 5+5, and urosternite VIII 1+1 macrosetae; all urosternal macrosetae are robust and with a few long barbs; robust smooth stylar setae. First urosternite appendages have (male) subcylindrical appendages thick, each of these has a large apical field with more than one hundred glandular a 1 setae., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 23, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Pacificampa wudonghuii Sendra sp. nov. urn:lsid:zoobank.org:act: 447EE93A-F466-4C66-827A-0751A259070F Figs 51–64 Etymology This species is dedicated to Professor Wu Donghui, from the Northeast Institute of Geography and Agroecology (NEIGAE), Chinese Academy of Sciences, Changchun (China), for his enthusiastic contribution and support to the knowledge of Northern China soil and cave biodiversity. Type material Holotype CHINA • ♀; Liaoning Province, Benxi, Huanren, Xianren Dong; 17 May 2019, L. Deharveng, A. Bedos and Wu Donghui leg.; labelled “CHILN19-005-holotype”; NEIGA-diplura-01. Paratypes CHINA • 1 ♀, 1 juvenile, same collection data as for holotype; labelled “CHILN19-005-female paratype”; MZB (MCNB) 2020-1160 • 1 juvenile; same collection data as for preceding; labelled “CHILN19- 005-juvenile paratype”; MZB (MCNB) 2020-1161 • 1 ♀; Liaoning Province, Benxi, Huanren, Pylon cave; 17 May 2019; L. Deharveng, A. Bedos and Wu Donghui leg.; labelled “CHILN19-007-female paratype”; MZB (MCNB) 2020-1164. Other material examined CHINA • 1 ♂, same collection data as for holotype; mounted on an aluminium stage and coated with palladium-gold; Coll. AS. Description BODY. Length 6.5–6.9 mm in females; 3.9 mm in one juvenile. Epicuticle smooth under optical microscope but well reticulated under high magnifications as one can see irregular polygonal structures of variable sizes with scattered external glands either visible or covered with secretion (Figs 51, 53); body with smooth clothing setae. HEAD. Antennae broken in all specimens; central antennomeres with two whorls of distal barbed macrosetae and uneven short setae; in addition, with single distal whorl of up to 8–12 gouge sensilla of 22–29 µm long (Fig. 51) and among them one or two small coniform sensilla (5 µm long). Proximal antennomeres with typical trichobothria disposition with bacilliform sensillum (9–10 µm long) on third antennomere in ventral position, between c–d macrosetae (Fig. 52). Plain frontal process with one anterior macrosetae, longer than clothing setae. Three macrosetae along each side of insertion line of antennomere and x setae with length ratios of a / i / p / x, 42/55/42/45, respectively, in holotype (Fig. 54). Large suboval labial palps, each with enlarged coniform latero-external sensillum near two gard setae and eight normal setae on anterior portion, up to 150 neuroglandular setae in medial and posterior positions. THORAX. Thoracic macrosetae distribution: pronotum has 1+1 ma, 1+1 la, 1+1 lp macrosetae; mesonotum has 1+1 ma, 1+1 la and 2+2 lp macrosetae; and metanotum has 1+1 ma and 1+1 lp macrosetae. Long macrosetae with long barbs in distal three-fourths; marginal setae longer than clothing setae, which are barbed from distal half to three-fourths. Legs elongated, metathoracic legs reach posterior border of seventh abdominal segment. Mesothoracic and metathoracic femora have one dorsal macroseta each, barbed along distal three-fourths (Fig. 55), absent in prothoracic femora. Calcars with two or three long barbs in middle (Fig. 56). Prothoracic and mesothoracic tibia with one short ventral macrosetae with one apical barb and two in metathoracic tibia (Fig. 57). Each tarsus with two separated ventral rows of thicker and longer setae among clothing setae, and a few setiform sensilla (Fig. 58). Three long smooth dorsal tarsal and one ventral setae. Subequal elbowed claws with smooth ventral surface ridged on dorsal side that can be mistaken for lateral crests under optical microscopes, between a blunt unguiculus and without lateral processes (Figs 59–60). ABDOMEN. Distribution of abdominal macrosetae on tergites (Fig. 61 shows 1+1 ma on III in the paratype from Grotte du Pylone, but absent in Xianren Dong Cave types; 1+1 ma, 1+1 la and 2+2 lp on IV–VII; 1+1 mp and 3+3 lp on VIII and 1+1 mp and 5+5 lp on IX abdominal; ma and la macrosetae with barbs in distal half to one third and shorter than mp and lp macrosetae, which bear long barbs along distal four-fifths. Urosternite I with 7+7 macrosetae (Fig. 62); urosternites II to VII with 4+4 macrosetae (Fig. 63); urosternite VIII with 1+1 macrosetae (Fig. 64); urosternal macrosetae of varying lengths, with one apical to twenty distal long barbs. Stylus setae smooth, apical and subapical setae shorter than ventromedial seta (Fig. 63). SECONDARY SEX CHARACTERS. Female urosternite I with subcylindrical appendages, each bearing up to 40 glandular a 1 setae in apical field. Male urosternite I with thick short, and subcylindrical appendages, each with large apical field of about 220 glandular a 1 setae (Fig. 62). Taxonomic affinities Pacificampa wudonghuii sp. nov. shares with P. birsteini Chevrizov, 1978, P. caesa Chevrizov, 1978 and P. nipponica Sendra, 2018 the same distribution of notal macrosetae (1+1 medial anterior, 1+1 lateral anterior, and 1+1 lateral posterior macrosetae on pronotum; 1+1 medial anterior, 1+1 lateral anterior, and 2+2 lateral posterior macrosetae on mesonotum; and 1+1 medial anterior and 1+1 lateral anterior macrosetae on metanotum). An undescribed species mentioned by Ferguson (1997) from northeastern China also has this setal distribution. However, P. wudonghuii sp. nov. differs from other species of Pacificampa in having 1+1 medial anterior, 1+1 lateral anterior, and 2+2 lateral posterior macrosetae on the fourth urotergite (Fig. 61), a pattern distribution that begins on the fifth urotergite for the other species. In addition, the larger male appendages have more abundant glandular a 1 setae in P. wudonghuii sp. nov. (Fig. 62) than in the other species of Pacificampa., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on pages 24-28, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205.","Sendra A., Yoshizawa K. & Ferreira R. L. 2018. New oversize troglobitic species of Campodeidae in Japan (Diplura). Subterranean Biology 27: 53 - 73. https: // doi. org / 10.3897 / subtbiol. 27.28575","Ferguson L. M. 1997. A report on a new species of Pacificampa (Diplura: Campodeidae) from a cave in China and a comparison of some North America genera to Pacificampa and Plutocampa previously only known from the Far East of Russia. Proceedings of the 12 th International Congress of Speleology La Chaux-de-Fonds, Neuchatel, Switzerland, August 10 - 17, Vol. 3: 315 - 317. Societe Suisse de Speleologie, Neuchatel."]}

Hubeicampa melissa Sendra & Lips gen. et sp. nov. urn:lsid:zoobank.org:act: E9EEC42A-21E5-43D8-B572-F4A6FD327323 Figs 20–48; Tables 3–4; Supp. file 3 Etymology This species is dedicated to the memory of Mélissa Lagrède, a French caver and friend of Josiane Lips. Mélissa died in an accident inside a nearby cave on the very day the holotype of this new species was collected. She also collected a paratype in the same cave the day before she died. Noun in apposition. Type material Holotype CHINA • ♀; Hubei, Banqiao, Grotte du 8 ième Ciel Baxian Dong; 8 Aug. 2009; Josiane Lips leg.; labelled “4168-holotype”; MZB (MCNB) 2020-1162. Paratypes CHINA • 1 ♂; same collection data as for holotype; labelled “4168-paratype”; MZB (MCNB) 2020-0621 • 1 ♀; same collection data as for holotype; Mélissa Lagrède leg.; 7 Aug. 2009; labelled “4158-paratype”; MZB (MCNB) 2020-1165. Other material examined CHINA • 2 specs; same collection data as for holotype; mounted on two separate aluminium stages and coated with palladium-gold; Coll. AS • 1 ♂; Sichuan, Beichuan, Yin Keng Dong; 22 Aug. 2011; Josiane Lips leg.; Coll. AS • 2 ♀♀; Hubei, Banqiao, Tie He Cave; 11 Aug. 2006; Josiane Lips leg.; Coll. AS. Description BODY. Length is 9 mm in holotype, 10.4 mm in female paratype and 6 mm in male paratype. Epicuticle smooth under optical microscope but reticulated at higher magnification, showing irregular polygonal structures of variable sizes (Figs 31–33). Body with apparently smooth clothing setae that in SEM have tiny expansions (micro–barbs) more abundant on other setae (macrosetae, marginal setae, style setae, tarsal setae and calcars) (Figs 26–29, 36, 42–45). HEAD. In holotype, one complete and intact antennae with 49 antennomeres; antennae slightly longer than body length (Table 3), with medial antennomeres 3 × as long as wide. Apical antennomere 4 × as long as wide, with irregular cupuliform organ containing ramiform, net-like complex with numerous perforations, occupying 1/10 of distal portion (Figs 20–21, 23). Distal and central antennomeres with two whorls of micro-barbed macrosetae (Figs 26–29); in addition, these have single distal whorl of up to 8–10 thin long gouge sensilla of 85–95 µm in length (Figs 22, 24), among them one small coniform sensillum (10 µm long). Proximal antennomeres with trichobothria on antennomeres 3–5 in a 2, 3, 1 pattern but with long flagellum (500–800 µm long in holotype). Third antennomere with bacilliform sensillum (25 µm long) in ventral position (Fig. 25). Coniform protrusion of frontal process with one anterior and four posterior micro–barbed macrosetae. Three macrosetae along each side of insertion line of antennomere and x setae with micro-barbs; ratios of length in female paratype a / i / p / x are 49/51/51/42, respectively. Suboval labial palps large, each with bacilliform latero-external sensillum near two simple setae, in posterior position with two guard setae and up to 24 setae on anterior portion, which are followed by up to 130 neuroglandular setae on posterior portion. THORAX. Distribution of thoracic macrosetae (Figs 30–33) with pronotum with 1+1 ma, 1+1 la 2 (1+2 la 2,3 in paratype), 2+2 lp 1,3 macrosetae; mesonotum with 1+1 ma, 1+1 la and 1+1 mp (0+0 mp in paratype) macrosetae, and metanotum with 1+1 ma, 0+1 smp (1+1 smp in paratype) macrosetae. All macrosetae slightly thin and micro-barbed, with a few longer and thin distal barbs; marginal setae longer than clothing setae. Legs elongated and metathoracic legs overpass end of abdomen. Tibia longer than femur or tarsus (Table 3). Femora III with one short dorsal macrosetae (0.17 mm in 1.88 mm femur of paratype-4158) (femora I–II with one dorsal macrosetae, less frequently without or with one submacrosetae instead). Calcar covered with thin abundant micro-barbs all over. Tibia I–III with one short completely micro-barbed ventral macroseta (in holotype, prothoracic legs of tibia with three and two macrosetae). Each tibia with two ventral rows of thick micro-barbed setae, scattered around tibia thin micro-barbed setiform sensilla. Three dorsal and one ventral tarsal subapical setae with micro-barbs (Figs 34, 36). Subequal elbowed claws with lateral crest; basal and ventral portions of claws covered with very small, thin, spiniform formations. Laminar lateral processes of the pretarsus completely covered with what was initially thought to be dense pubescence, but under SEM high magnifications turned out to be thick micro-barbs (Figs 34, 36–39). ABDOMEN. Distribution of abdominal macrosetae on tergites (Fig. 40): 1+1 post 1 on V tergite, (1 post 1 + 2 post 1,5 on paratype); 3+3 post 2,4,5 on VI tergite, 3+3 post 1,4,5 (3+2 post 4,5 in holotype) on VII tergite, 4 post 1,3,4,5 + 5 post 1–5 (5+5 (post 1–5 in paratype) on VIII tergite; and 6+6 post on IX abdominal segment. Short micro-barbed abdominal macrosetae with few distal barbs on urotergites V– VII; urotergite VIII and abdominal segment with similar but longer macrosetae. Urosternite I with 12+12 macrosetae (Fig. 41); urosternites II to VII with 5+5 macrosetae; urosternite VIII with 1+1 macrosetae; urosternal macrosetae complete covered by micro-barbs including a few distal short barbs. Stylus with apical, subapical and ventromedial setae completely covered with both micro-barbs and one apical barb (Figs 42–43). Cerci 1.7 × as long as body length in two presumably regenerated cerci of paratype, with six primary articles (Table 4). Length of cerci quickly increases from proximal to distal articles, covered with long setae, and macrosetae covered with micro-barbs; each primary article with single apical whorl of shorter and thinner setae also covered with micro-barbs (Figs 44–45). SECONDARY SEX CHARACTERS. Female urosternite I with coniform appendages, each bearing up to 12 glandular a 1 setae in apical field (Fig. 41). Male urosternite I with subcylindrical enlarged appendages with large field of glandular a 1 sertae; without glandular g 1 setae.

Genus Plutocampa Chevrizov, 1978 Diagnosis (modified and improved from the original diagnosis by Chevrizov 1978) Thick sensilla on third antennomere and labial palps. Thoracic macrosetae with no less than 5+5 macrosetae on pronotum, 5+5 on mesonotum (1+1 medial posterior included), and 4+4 on metanotum (1+1 medial posterior included). Two dorsal macrosetae on metathoracic femora and one on tibia. Unequal elbowed claws with lateral crests without lateral processes. Urotergites V–VII with 4+4 posterior macrosetae and without lateral anterior macrosetae. Up to 6+6 macrosetae on first urosternite, 4+4 macrosetae on second to seventh urosternites, and 1+1 macrosetae on eighth urosternite., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 20, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Pacificampa birsteini Chevrizov, 1978 Pacificampa birsteini Chevrizov, 1978: 198, fig. 1(1‾9). Material examined RUSSIA • ♀ holotype (7 mm); Far East, Primoriye region, Partizanskii Mountain Range, Belyi Dvorets Cave (White Palace Cave); 16 Aug. 1966; S.I. Ljovushkin leg.; slide; ZMMU Dip0003. Redescription (based on Chevrizov 1978) It has a smooth cuticle, under optical microscope, covered by abundant short smooth clothing setae. The antennae are apparently intact with 26 antennomeres; the medial antennomeres are 3 × as long as wide, and the last antennomere has about eight spheroidal olfactory chemoreceptors; the third antennomere has a sensillum in ventral position (c–d macrosetae). The plain frontal process has one long anterior smooth macrosetae and two, almost two times shorter, posterior macrosetae; the three macrosetae have a few thin distal barbs along each side of the insertion line of antennomere longer than the smooth x setae, with length ratios of a / i / p / x, 5/6/5/2, respectively. Pronotum with 1+1 ma, 1+1 la, 1+1 lp, mesonotum with 1+1 ma, 1+1 la, 2+2 lp, and metanotum with 1+1 ma, 1+1 lp macrosetae; all notal macrosetae are robust with a few short distal barbs; the marginal setae are thicker than the clothing setae with short distal barbs, whereas the lateral posterior marginal setae are larger with abundant small barbs. The metathoracic legs overpass the end of the abdomen; the femora have one dorsal robust macroseta with 2–3 distal barbs, the tibia has two short ventral macrosetae with 1–2 apical barbs and the calcars have 3–4 long barbs; the tarsus has two rows of smooth robust ventral setae, three long smooth dorsal and one distal ventral setae; the pretarsus is without lateral processes, with subequal elbowed claws and with a smooth ventral surface, ridged on the dorsal side, that looks like very small lateral crests under optical microscope, but they are not. Urotergite IV with 1+0 la, 0–1 lp; urotergites V–VII with 1+1 ma, 1+1 la, 2+2 lp; urotergite VIII and abdominal segment IX with 1+1 mp, 3+3 lp, all macrosetae are robust with a few short distal barbs. Urosternite I with 6+6, urosternites II–VII with 5+5, and urosternite VIII with 1+1 macrosetae; all urosternal macrosetae are robust with a few long barbs; robust smooth stylar setae. First urosternite appendages (female) are subcylindrical and long with an apical field of a 1 glandular setae. Cercal articles have whorls of long macrosetae with a few distal barbs., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on pages 22-23, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Plutocampa paurochaeta Chevrizov, 1978 Plutocampa paurochaeta Chevrizov, 1978: 204, fig. 3(11‾19). Material examined RUSSIA • ♀ holotype (6.7 mm); Far East, Primoriye region, Chandolaz Mt. Range, Solyanik Cave; 25 Jun. 1974; P.B. Chevrizov leg.; slide; ZMMU Dip0005. Redescription (based on Chevrizov 1978) Under optical microscope, one can see a smooth cuticle covered by thin smooth clothing setae. Antennae, apparently intact, have 28 antennomeres, the central antennomere is 1.6 × as long as wide, the apical antennomere has about eight olfactory chemoreceptors; the third antennomere has a large sensillum in ventral position. The head has a small and pointed frontal process with one long anterior smooth macroseta and two shorter posterior macrosetae; three smooth macrosetae along each side of the insertion line of the antennomere and smooth x setae with similar length, ratios of a / i / p / x are 6/5/8/7, respectively. Pronotum with 1+1 ma, 2+2 la, 2+2 lp 1,3 , mesonotum with 1+1 ma, 2+1 la, 2+2 lp 2,3 , and metanotum with 1+1 ma, 2+2 lp macrosetae, notal macrosetae are long and thin with thin barbs on three-fourths of distal barbs; marginal setae are long and smooth. Metathoracic legs overpass the end of the abdomen; femora have two long barbed dorsal macrosetae and one ventral macroseta, and the tibia has one ventral short macroseta and calcars with several long barbs. The tarsus has two rows of ventral smooth setae slightly thicker than the clothing setae, three long and smooth dorsal subapical setae and one ventral subapical seta; the pretarsus is without lateral processes, with unequal elbowed claws, and with large lateral crests. Urotergite IV with 1+1 post 1 , urotergites V–VII with 4+4 post 1–4 , urotegite VIII with 5+5 post1–5, macrosetae are long, with barbs along their three-fourths. Urosternite I with 6+6, urosternites II–VII with 4+4, and urosternite VIII with 1+1 macrosetae; urosternal macrosetae are shorter than urotergal macrosetae with long barbs; smooth stylar setae. First urosternite appendages (female) have large, short subconical glandular a 1 setae with a smaller apical field. The cerci have eight articles plus a basal one covered with whorls of smooth setae and whorls of macrosetae barbed with thin barbs at the proximal articles to smooth in medial and distal articles., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 22, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Chevrizov B. P. 1978. Two new genera of the Family Campodeidae from the Far East Caves. Zoologichesky Zhurnal 57 (2): 197 ‾ 205."]}

Genus Anisuracampa Xie & Yang, 1991 Diagnosis (modified and improved from original diagnosis by Xie & Yang 1991) Mesonotum and metanotum with 1+1 medial anterior, no or 2+2 lateral anterior, and 2+2 to 4+4 lateral posterior macrosetae. Two to three dorsal macrosetae on metathoracic legs. Elbowed claws ventrally with long spiniform formations with apparently lateral crests. Laminar pretarsus lateral processes with long, broad barbs. Urotergites with 4+4–5+5 posterior macrosetae on urotergites V–VII. Eighth urosternite with 1+1 macrosetae. First urosternite with non–glandular setae bearing coniform or subtrapezoidal appendages., Published as part of Sendra, Alberto, Komerički, Ana, Lips, Josiane, Luan, Yunxia, Selfa, Jesús & Jiménez-Valverde, Alberto, 2021, Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha), pp. 1-46 in European Journal of Taxonomy 731 on page 3, DOI: 10.5852/ejt.2021.731.1199, http://zenodo.org/record/4422557, {"references":["Xie R. & Yang Y. 1991. Description of two new genera and three new species of Campodeidae in China (Diplura). Contribution Shanghai Institute of Entomology 10: 95 ‾ 102."]}

Campodea (Paurocampa) rocasolanoi Silvestri, 1932 Material studied: Pre-Pyrenees: Sierra de Santo Domingo, Umbr��a de Raya Alta; 2 ♂♂, 1 ♀, scree slope MSS, 1 SSD, 2 IX 2015. V.M. Ortu��o, J.D. Gilgado, E. Cuesta & E. Ledesma leg. Sierra de Santo Domingo, Solana del Portillo de Long��s; 1 ♂, 7♀♀, scree slope MSS, 3SSD, 2 IX 2015. 1 ♀, scree slope MSS, 4SSD, 2 IX 2015. V.M. Ortu��o, J.D. Gilgado, E. Cuesta & E. Ledesma leg. Habitat and distribution: It is spread along the west of Pyrenees and the Cantabrian regions, dwelling in the upper layer of moist soils., Published as part of Sendra, Alberto, Jim��nez-Valverde, Alberto, Gilgado, Jos�� D., Ledesma, Enrique, Baquero, Enrique, P��rez-Su��rez, Gonzalo, Cuesta, Eva, Herrero-Borgo����n, Juan J., Jordana, Rafael, Tinaut, Alberto, Barranco, Pablo & Ortu��o, Vicente M., 2017, Diplurans of subsurface terrestrial habitats in the Iberian Peninsula, with a new species description (Diplura: Campodeidae), pp. 61-80 in Zootaxa 4291 (1) on page 72, DOI: 10.11646/zootaxa.4291.1.4, http://zenodo.org/record/829369, {"references":["Silvestri, F. (1932) Campodeidae (Thysanura) de Espana (primera parte). Eos, 8, 115 - 164."]}

Campodea (Dicampa) neusae Sendra & Moreno, 2006 Material studied: Central System Mountains: Guadarrama National Park-Arroyo del Charco, 2 ♀♀, scree slope MSS, 25 SSD, 22 X 2015, GIBSES leg., 1 ♂, 4 ♀♀, scree slope MSS, 25SSD, 9 VI 2016. GIBSES leg. Guadarrama National Park, Navafría Mountain Pass, 1 ♂, scree slope MSS in pine forest, 13 VI 2016. GIBSES leg. Morphological remarks. One specimen has an intact antenna with 24 antennomeres. Habitat and distribution. This species was found in a single location in the Natural Park Tejera Negra in the eastern extreme of the Central Systems Mountains among leaves in forest soil.

Campodea (Campodea) fragilis Meinert, 1865 Material studied. Prebaetic Mountains: Xal�� river; 1 ♂; alluvial MSS, 2 SSD, 8 III 2012; 1 specimen, sex unknown, alluvial MSS, 2SSD, VII 2012. GIBSES leg. Habitat and distribution. This is perhaps the most widespread species of dipluran occurring in much of the Holarctic region. It is an epiedaphic species frequently found near the water table, rivers or lakes. Within the Iberian Peninsula it has not yet been found along the Atlantic coast (Sendra & Moreno 2004)., Published as part of Sendra, Alberto, Jim��nez-Valverde, Alberto, Gilgado, Jos�� D., Ledesma, Enrique, Baquero, Enrique, P��rez-Su��rez, Gonzalo, Cuesta, Eva, Herrero-Borgo����n, Juan J., Jordana, Rafael, Tinaut, Alberto, Barranco, Pablo & Ortu��o, Vicente M., 2017, Diplurans of subsurface terrestrial habitats in the Iberian Peninsula, with a new species description (Diplura: Campodeidae), pp. 61-80 in Zootaxa 4291 (1) on page 69, DOI: 10.11646/zootaxa.4291.1.4, http://zenodo.org/record/829369, {"references":["Sendra, A. & Moreno, A. (2004) El subgenero Campodea s. str. en la Peninsula Iberica (Hexapoda: Diplura: Campodeidae). Boletin de la Sociedad Entomologica Aragonesa, 35, 19 - 38."]}

Campodea (Campodea) posterior Silvestri, 1932 Material studied. Penibaetic Mountains: Cirque of Veleta summit; 1 ♀, 3 juveniles, scree slope MSS, 3 SSD, IX 2012, 2 ♀♀, scree slope MSS, 3SSD, X 2015. V.M. Ortuño, J.D. Gilgado & A. Tinaut leg. Habitat and distribution. Endemic to the Iberian Peninsula, C. posterior is an abundant species spread along the Baetic Mountains living in both humid and dry soils at various depths (Sendra & Moreno 2004).

Sendra, Alberto, Jiménez-Valverde, Alberto, Gilgado, José D., Ledesma, Enrique, Baquero, Enrique, Pérez-Suárez, Gonzalo, Cuesta, Eva, Herrero-Borgoñón, Juan J., Jordana, Rafael, Tinaut, Alberto, Barranco, Pablo, Ortuño, Vicente M. (2017): Diplurans of subsurface terrestrial habitats in the Iberian Peninsula, with a new species description (Diplura: Campodeidae). Zootaxa 4291 (1): 61-80, DOI: https://doi.org/10.11646/zootaxa.4291.1.4

Campodea (Dicampa) escalerai Silvestri, 1932 Material studied. Iberic System Mountains: Moncayo Massif, Barranco de Castilla; 1 juvenile, scree slope MSS in beech forest, 2 SSD, 11 VII 2013. 1 ♀, scree slope MSS in a beech forest, 2SSD, 25 IX 2012. V.M. Ortu��o, J.D. Gilgado, E. Cuesta & E. Ledesma leg. Habitat and distribution. Campodea escalerai shares habitat and distribution with C. (D.) catalana, but occurs in a smaller geographical area and is much less abundant (Sendra & Moreno 2006)., Published as part of Sendra, Alberto, Jim��nez-Valverde, Alberto, Gilgado, Jos�� D., Ledesma, Enrique, Baquero, Enrique, P��rez-Su��rez, Gonzalo, Cuesta, Eva, Herrero-Borgo����n, Juan J., Jordana, Rafael, Tinaut, Alberto, Barranco, Pablo & Ortu��o, Vicente M., 2017, Diplurans of subsurface terrestrial habitats in the Iberian Peninsula, with a new species description (Diplura: Campodeidae), pp. 61-80 in Zootaxa 4291 (1) on page 72, DOI: 10.11646/zootaxa.4291.1.4, http://zenodo.org/record/829369, {"references":["Silvestri, F. (1932) Campodeidae (Thysanura) de Espana (primera parte). Eos, 8, 115 - 164.","Sendra, A. & Moreno, A. (2006) Los subgeneros Dicampa, Monocampa y Paurocampa en la Peninsula Iberica e Islas Baleares (Diplura: Campodeidae). Boletin de la Sociedad Entomologica Aragonesa, 39, 69 - 82."]}

Campodea (Campodea) portacoeliensis Sendra & Jiménez, 1986 Material studied. Central System Mountains, Miraflores de la Sierra, near to Morcuera Mountain Pass; 1 ♂, scree slope MSS Reajo Cazado Stream, 13 VI 2016. GIBSES leg. Habitat and distribution. This epiedaphic species founds in the uppermost soil layer between the leaves. It is distributed in humid spots in the Iberic Mountains, Eastern of the Iberian Peninsula (Sendra & Jiménez 1986; Sendra 1989b; Sendra & Moreno 2004).