Asian cave-adapted diplurans, with the description of two new genera and four new species (Arthropoda, Hexapoda, Entognatha)

Twenty-nine specimens of Diplura collected from eight caves in China and Myanmar contain two new genera, Hubeicampa Sendra & Lips gen. nov. and Mueggejapyx Sendra & Komerički gen. nov., as well as four new species, Anisuracampa ywangana Sendra & Komerički sp. nov., Hubeicampa melissa Sendra & Lips gen. et sp. nov., Pacificampa wudonghuii Sendra sp. nov. and Mueggejapyx brehieri Sendra & Komerički gen. et sp. nov. These cave-adapted taxa showcase an impressive diversity in morphological adaptation (troglomorphy) to cave ecosystems. Their sensorial equipment, setae and receptors in the cupuliform organ have unique forms (H. melissa gen. et sp. nov.), as well as the pretarsus sticky surface (A. ywangana sp. nov. and H. melissa gen. et sp. nov.). Recent contributions European Journal of Taxonomy 731: 1–46 (2021) 2 on Asian diplurans, together with the taxonomic novelties shown in the present study, highlight the biogeographical importance of the Asian biodiversity. Asia is revealed as a continent with vast karst regions still waiting to be explored and new dipluran species waiting to be discovered.


Introduction
Asia is the largest continent and has 30% of the Earth's total land surface, in which karst landscapes with caves occupy vast regions; two million square kilometers in China (Juberthie & Decu 2001). A huge number of these caves remain unexplored from a biospeleological perspective (Juberthie & Decu 2001). Over the last two decades, studies of Asian caves have revealed a wide diversity of highly caveadapted species in Collembola, Carabidae, Diplopoda, fishes, and other zoological taxa (Ma & Zhao

Material and methods
Specimens were collected directly by hand and stored in 70-75% ethanol, then washed with distilled water and mounted on slides in Marc André II solution. Mounted specimens were examined with a phase-contrast optical microscope (Leica DMLS). The illustrations were made with a drawing tube and measurements were taken with an ocular micrometer. For measuring the body length, the specimens were mounted in toto and measured from the base of the frontal process distal macrosetae to the abdomen's supra-anal valve. Two specimens, coated with palladium-gold, were used for SEM (Hitachi S-4800) photography and for measuring the sensilla.
Terminology for the species is from Smith (1962) and Pagés (1953Pagés ( , 1984: A = macrosetae in A position; B = macrosetae in B position; C = macrosetae in C position; D = macrosetae in D position; L = macrosetae in L position; M = larger macrosetae of the body set in reinforced setal sockets; ms = microsetae or minute setae visible only under high magnification and always set in simple setal sockets; P = macrosetae in P position; sM = medium-sized sub-macrosetae usually set in simple setal sockets; V = macrosetae in V position; friction setae = microsetae with large sockets that occurs in groups where the body integument folds or moves upon itself and would perhaps otherwise cause abrasion. For the specific terminology of buccal pieces see Pagés (1964) and for thoracic sclerites see Pagés (1964) and Barlet & Carpentier (1962). For the cercal chaetotaxy we have used Muegge & Bernard (1990).
The holotype, paratypes and other studied specimens are kept at the following institutions:

Anisuracampa ywangana
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 . 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  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 lp2,3 macrosetae; mesonotum with 1+1 ma, 2+2 la and 4+4 lp1-4 macrosetae; and metanotum with 1+1 ma, 2+2 la and 3+3 lp1-3 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 post1 on I-III; 1+1 post1 to 3+3 post1-3 on IV, 4+4 post1-4 and 1+1 la on V, 5+5 post1-5 or 6+6 post1-6 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 . 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 a2 setae proximally and apical field of up to 28 glandular a1 setae . Male with similar appendages with two fields of glandular setae (19 a2 and 27 a1) 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. 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

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.

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 . 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) 36,(42)(43)(44)(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 23). Distal and central antennomeres with two whorls of micro-barbed macrosetae ; 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.
Secondary Sex cHaracTerS. Female urosternite I with coniform appendages, each bearing up to 12 glandular a1 setae in apical field (Fig. 41). Male urosternite I with subcylindrical enlarged appendages with large field of glandular a1 sertae; without glandular g1 setae.

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.

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. Chevrizov, 1978 Figs 49-50 Plutocampa methoria Chevrizov, 1978: 202, fig. 3(1−10). Redescription (based on Chevrizov 1978) Under optical microscope, one can see a smooth cuticle covered by thin and smooth clothing setae. The more complete antennae is apparently regenerated and it has 23 antennomeres in the 5.4 mm adult from Grotte du Pylone; the third antennomere has a thick sensillum in ventral position (d-e macrosetae) (Fig. 49). The small and pointed frontal process has one long and smooth anterior macroseta and two shorter posterior macrosetae; there are 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 14/19/14/19 in the holotype, respectively. The labial palps have a large thick sensilla (Fig. 50). Pronotum with 1+1 ma, 2+2 la, 2+2 lp 1,3 , mesonotum with 1+1 ma, 2+2 (2+1) la, 2+2 lp 2,3 , 1+1 mp, and metanotum with 1+1 ma, 0-1+1 la, 2+2 lp 2,3 , 1+1 mp macrosetae; long and thin notal macrosetae with thin barbs along their distal three-fourths; marginal setae are long and barbed as the macrosetae. Metathoracic legs reach the posterior border of the V-VI abdominal segment; femora has two long, barbed dorsal macrosetae and one ventral macroseta, while the tibia has one short ventral macroseta, a few distal barbs, and calcars with several long barbs; the tarsus has two rows of smooth ventral setae slightly thicker than the clothing setae, and in the distal portion: three long, smooth dorsal setae, one ventral seta and one distal seta; the pretarsus is without lateral processes, with unequal elbowed claws and large lateral crests. Urotergite I-II with 1+1 post 1 ; urotergite III with 2+2 post 1,2 ; urotergites IV-VII with 4+4 post 1-4 ; urotegite VIII with 5+5 post 1-5 and abdominal segment IX 7+7 with post, macrosetae are long with barbs along their distal three-fourths. Urosternite I with 6+6, urosternites II-VII with 4+4 and urosternite VIII with 1+1 macrosetae; urosternal macrosetae shorter than urotergal macrosetae with long barbs; smooth stylar setae. Appendages of the first urosternite (female) with large, short and subcylindrical appendages, each with apical field of glandular a 1 setae. The most complete cercus is regenerated and it has six primary articles and a basal article, which are covered by several whorls of barbed macrosetae with long barbs along the third distal and a few whorls of smooth setae.

Material
Figs 49-50. Plutocampa methoria Chevrizov, 1978. 49. Sensillum of the third antennomere. 50. Left labial palp and palpiform process (pp). Chevrizov, 1978 Plutocampa paurochaeta Chevrizov, 1978: 204, fig. 3(11−19). 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 post 1-5 , 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.

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. Chevrizov, 1978 Pacificampa caesa Chevrizov, 1978: 199, figs 1(10−19).

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, 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).

Taxonomic notes
The distribution of notal and urotergal macrosetae on the immature male is the same as that of P. birsteini Chevrizov, 1978 andP. caesa Chevrizov, 1978

Diagnosis
Large and elongated body; antennae with 36 antennomeres; trichobothria on antennomeres 4-6 in a 3/5/5 pattern; a trichobothria in distal position; 1-2 placoid sensilla on medial and distal antennomere; apical antennomere with 16-18 placoid sensilla; dorsal and ventral side of head with numerous uniformly distributed sM and sm, and several M; five laminae pectinate on maxilla; elongated thoracic segments; and cuticle with tiny micro-holes, pronotum and mesonotum each with 5+5 M, metanotum with 2+2 M; strong internal Y cuticular furcisternite structures in pro-presternites and pro-, meso-and metasternites; urotergites with at most 1+1 M (ma), 1+1 M 1 , 1+1 M 2-5 ; urite X longer than wide, with marked carinae with subparallel margins, with 2+2 intracarinal macrosetae; acropygium rounded; lateral angles of urotergites VI and VII with weakly to moderate lobiform projection; first urosternite with multiperforated surface and rounded protrusions, with 3+3 M on prescutum and 11+11 M on scutum; median glandular organ with seta-shaped sensilla; lateral subcoxal organs with one to three rows of short glandular setae (GS), one row of sensory setae (SS), and a row of setae with large socket; sternites with abundant sm and strong M; cerci strong, elongated, rectilinear and becoming curved subapically, heavily sclerotized with external dorsal and ventral carinae. Both cerci without medial tooth, but with one short row of small round denticles on the right cercus and two rows of small round denticles on the left cercus.

Etymology
This genus is dedicated to the memory of the American entomologist Mark Alan Muegge (pronounced Meggy), who died young due to an unfortunate car accident in 2015. He left behind a promising career and was devoted, among other groups, to diplurans; alongside other taxa, he provided a description of the highly cave-adapted japygid from Texas, Mixojapyx reddelli Muegge, 1992. Sendra & Komerički gen. et sp. nov. urn:lsid:zoobank Paratypes MYANMAR • 1 ♂, 1 ♀; same collection data as for holotype; labelled "♂3-paratype" and "♀1- Other material examined MYANMAR • 2 specs; same collection data as for holotype; mounted on two separate aluminium stages and coated with palladium-gold; Coll. AS.

Taxonomic affinities
Mueggejapyx gen. nov. is distinguished by its elongated, rectilinear, heavily sclerotized cerci curving subapically, and lacking a medial tooth but with one or two rows of basal small round denticles, its median glandular organ with seta-shaped sensilla, and short lateral subcoxal organs, each with a row of setae that have large sockets.

Remarks
Mueggejapyx brehieri gen. et sp. nov., was observed and collected along with A. ywangana sp. nov. in Win Twin Cave (Figs 1-4), as well as in two nearby caves: Parpent Cave 1 and Kyauk Khaung (synonyms: Sin Lae Ye Win, Stone Cave). The latter is the second longest cave so far explored in Myanmar. Kyauk Khaung Cave is over 4790 m long and also situated in the Ywangan karst area. Its entrance is located only 1240 m southeast of the entrance to Win Twin, and the cave itself extends mostly towards the northeast of the entrance. During the 2015 biological research of Kyauk Khaung, specimens of Campodeidae and Japygidae were found in the northernmost parts of the cave, which indicates the species is most likely distributed throughout the entire subterranean habitats of the Ywangan karst (unpublished data). The Ywangan karst covers 1050 ha, and at the moment hosts three other endemic troglobitic invertebrates: the carabid beetle Birmaphaenops brehieri Deuve, 2017(Deuve 2017, the cave crab Shanphusa ywarngan Ng & Whitten, 2017(Ng & Whitten 2017, and a woodlouse Bamaoniscus lobatus Taiti et al, 2020 as well as an endemic karst-adapted gecko, the Linn-Way bent-toed gecko Cyrtodactylus linnwayensis Grismer et al., 2017(Grismer et al. 2017, and 17 species of bats (unpublished data). Due to its biodiversity, Ywangan karst is proposed as a karst Key Biodiversity Area and a subterranean
Campodeinae has only one cave genus, Pacificampa, with five species: two in the extreme east of Siberia (Chevrizov 1978); two on the Japanese islands of Kyushu and Shikoku (Sendra et al. 2018) and P. wudonghuii sp. nov. from northeastern China. All five species have in common their peculiar enlarged dorsal ridged claws without pretarsus lateral processes (Figs 59-60), similarities in the distribution of the macrosetae in the thorax and abdomen, as well as the absence in both sexes of glandular g 1 setae on the posterior border of the first urosternite and glandular a 2 setae on the first urosternite appendages (Fig. 62).
Plusiocampinae is diverse and widespread in cave areas of Asia with eight genera in Asian caves: Anisuracampa with one cave species in Myanmar and another in soil in China; Plusiocampa with one cave species in the Anatolian peninsula, another in the western Caucasus and numerous cave and soil species limited to the Euro-Mediterranean region; Plutocampa with three species in the Russian Far East and northeast China; Hubeicampa gen. nov. with two species in central China; and five monotypic genera: Vandelicampa Condé, 1955 in the most western part of Asia, Simlacampa Condé, 1956 in northwest India, Turkmenocampa in central Asia, and Whittencampa in southern China. All these genera show important differences in their pretarsal structures, with unclear relationships between them. Pretarsus lateral crests of claws with different sizes and ornamentation are present in all genera except Turkmenicampa, which has a lateral, sharp side-piece on each claw. However, the pretarsus lateral processes vary among the genera: setiform (Plusiocampa), subcylindrical (Whittencampa) or absent (Plutocampa), and laminar in the rest of the genera. The laminar lateral processes are long and narrow (Simlacampa and Anisuracampa), short and spiniform (Hubeicampa gen. nov.), or long, spiniform (Vandelicampa) (Condé 1955(Condé , 1956(Condé , 1993Chevrizov 1978;Sendra et al. 2017a, Sendra & Deharveng 2020. Simlacampa and Anisuracampa have a very similar pretarsus (claws with lateral crests, and laminar and divided lateral processes). The complete pretarsus of Simlacampa was drawn with full details by Condé (1956: figs 1-2), allowing it to be compared with SEM images of Anisuracampa ywangana sp. nov. (Figs 12-15). However, at least two differences separate these genera. Anisuracampa has two or three dorsal macrosetae on the metathoracic femur compared to none in Simlacampa. Also, Anisuracampa has numerous macrosetae on the nota (Fig. 9) and urotergites (medial anterior, lateral anterior, and lateral posterior macrosetae on the meso and metanotum, 4+4-5+5 posterior macrosetae on the urotergites V-VII) compared with less macrosetae in Simlacampa (l+1 lateral posterior macrosetae on the mesonotum and none on the metanotum; 2+2 posterior macrosetae on the urotergite VII). The pretarsus in other genera of Plusiocampinae has some similarities with that of Anisuracampa and Simlacampa, such as Cestocampa Condé, 1956;Patrizicampa Condé, 1956 or Vandelicampa, with broad laminar pretarsal lateral processes (Sendra et al. 2017), but closer study needs to be made with SEM or high-quality optical microscopy.
Hubeicampa gen. nov. proposed for Hubeicampa melissa gen. et sp. nov. and Hubeicampa lipsae (Condé 1993) comb. nov. (= Plusiocampa (Dydimocampa) lipsae Condé, 1993), inhabiting caves in Hubei, central China, is characterized by a unique pretarsus with subequal, elbowed claws and lateral crests as in many Plusiocampinae, except that the basal and ventral portions of the claws are covered with numerous, very small spiniform formations that appear pubescent under an optical microscope, and the laminar pretarsal lateral processes, which are completely covered with dense micro-barbs (Figs 36-39).
In addition, species of Hubeicampa gen. nov. are characterized by a singular kind of pubescent setae and macrosetae covered with micro-barbs visible under high magnifications (Figs 26-29).
Cave-adapted japygids are taxonomically defined, due to their genetic isolation, and thus their genera are well settled and separated from other taxa (Pagés 1980;Muegge 1992;Sendra et al. 2006b). In continental Asia, cave-adapted species are represented by Kohjapyx lindbergei Pagés, 1962 from a cave in Afghanistan, Burmjapyx inferus (Carpenter, 1932) from Batu Caves in Malaysia (Pagés 1962(Pagés , 1964 and Mueggejapyx brehieri gen. et sp. nov. from four caves in western Myanmar. These taxa especially differ in cercal shape and distribution patterns of teeth and denticles. The loss of the medial cercal tooth is the most distinguishing feature of Mueggejapyx gen. nov., which is only shared with three other genera: Psalidojapyx Pagés, 2000 with two species from Borneo; Rectojapyx Pagés, 1953 with one species from Europe and Rossjapyx Smith, 1962 with two species from Chile, all of which are soil-dwelling (Pagés 1953(Pagés , 2000Smith 1962). Mueggejapyx gen. nov. differs in the unique shape of its cerci, which are strong and well developed, elongated and rectilinear, becoming curved subapically. Each cercus has exterior dorsal and ventral carinae and very short rows of small basal denticles preceded by a basal notch on the right cercus and a round tooth on the left cercus (Figs 91-95). Another important feature of Mueggejapyx gen. nov. is the presence of mostly shorter lateral subcoxal organs occupying less than ⅓ of the interstyle width with smaller glandular setae. Cave-adapted features of M. brehieri gen. et sp. nov. include extra placoid sensilla on the antennomeres from the middle of the antennae to the apex, with numerous placoid sensilla on the apical antennomere (Figs 65-66), and a substantial lengthening of thoracic segments and body in general (Figs 71,75).

Troglomorphy
The handful of cave-adapted campodeid species already known in Asia show highly adapted features to cave ecosystems. In campodeids, the longest bodies with elongated segments, particularly of the thorax, belong to Pacificampa daudarabochi Sendra, 2018 and Hubeicampa melissa gen. et sp. nov., being more than one centimetre in length. In all Asian cave species, the antennae, legs and cerci are elongated: metathoracic legs reach or overpass the end of the abdomen; the antennae are as long as or longer than the body length; and the cerci can be twice the body size in some species (e.g., Whittencampa troglobia Sendra & Deharveng, 2020) (Sendra & Deharveng 2020). The elongation of appendages in cave-adapted species is due to the increase in article length and/or more numerous antennomeres and cercal articles (Sendra et al. 2017b). The lengths of the middle antennomere are twice that of the width and the length of the apical antennomere is four times that of the width in species of Hubeicampa gen. nov., P. daudarabochi and W. troglobia. However, in cave-adapted species of Lepidocampinae, the antennomere length and width are equal but the antennomeres increase in number (Sendra et al. 2017b).
The antennomeres and cercal articles in Asian cave-adapted species can reach very high numbers, e.g., in W. troglobia with 54 antennomeres, the highest in Plusiocampinae (Sendra & Deharveng 2020 Sendra & Stoev, 2017) or caliciform shape with folds in radial expansions from a central axis (P. daidarabotchi) (Sendra et al. 2017(Sendra et al. , 2018. The mechanoreceptor setae reach a unique shape with tiny expansions called micro-barbs in the highly cave-adapted H. melissa gen. et sp. nov., which are not seen in other campodeids (Figs 26-29).
The pretarsal structure adaptations in Plusiocampinae show an important variation range, apparently correlated with cave-habitat. Cave-dwelling species of Plusiocampa have larger claws and bigger lateral crests than their soil relatives (Condé 1956;Sendra et al. 2020). The claws have lateral crests, except for T. mirabilis which has two sharp side-piece expansions instead. The laminar lateral processes of Anisuracampa, Hubeicampa gen. nov., and Turkmenocampa each have narrow laminar expansions (Figs 12-15), very small barbs (Figs 36-39), or long barbs (Sendra et al. 2017a) with more or less apical enlargements that seem to have sticky properties, allowing those diplurans to walk on smooth surfaces (e.g., speleothems).
The three exclusively Asian cave-adapted japygids show different stages of troglomorphy. Burmajapyx inferus shows no elongation in body or appendages, and although it has extra placoid sensilla on the medial and distal antennomeres, it does not have extra sensilla on the apical antennomere. Kohjapyx lindbergi Pagés, 1962 shows a moderate elongation in body and appendages and has extra placoid sensilla on the apical antennomere. Mueggejapyx brehieri gen. et sp. nov. is noticeably elongated (Figs 2, 71, 75), especially in the thoracic segments, and has extra placoid sensilla on the medial, distal, and apical antennomeres (Figs 65-66).

Diversity and distribution
Although there are several studies dealing with local Asian dipluran faunas (e.g., Silvestri 1931Silvestri , 1948Chou 1983Chou , 1984Pagés 1984Pagés , 2002Condé 1990;Sendra et al. 2006aSendra et al. , 2010; among others), the Asian diplurans have never been the subject of a comprehensive work. A total of 197 species of diplurans has been described or cited from soil and cave habitats from 508 localities, mostly distributed throughout western and eastern or southeastern Asia. The 21 known Asian cave-adapted dipluran species follow this same general distribution pattern, although they are unavoidably restricted to the vast karst regions of Asia (Figs 98-99; Supp. file 1). Nevertheless, cave diplurans have not been found in desert zones, probably due to the lack of infiltration of organic matter into cave ecosystems (Sendra & Reboleira 2014) (Figs 98-99). Furthermore, diplurans cannot enter hypogenic cave systems, although they can thrive in there after the erosion of the confining rock layers (Klimchouk 2007;Jimenez-Valverde et al. 2017) (Figs 98-99). A very important historical factor that has conditioned the current distribution of cave diplurans is the former extension of glacial and permafrost areas during the Last Glacial Maximum (Fig. 98). These frozen conditions presumably eliminated terrestrial cave animals from vast geographical regions that have not yet been recolonized, something that has been shown for other taxonomic groups with limited dispersal abilities (e.g., Svenning & Skov 2007). Finally, although there are many vast karst areas in China without records, these are covered, even if in a patched way, by permafrost at present, so the presence of cave-adapted diplurans seems unlikely (Fig. 99). In contrast, there are extensive karstic areas in southeast China that have the potential to harbour new species to science (Figs 98-99).
The different taxonomic groups of Asian cave-adapted diplurans have an unequal representation in the different Asian regions. While japygids seem to barely colonize cave-ecosystems, with three currently known species belonging to three genera, campodeids have a much larger presence in caves, with 18 already described species belonging to 10 genera from western to eastern Asian regions. Within  Condé, 1956 (brown circles), Japygidae Haliday, 1864 (red squares). In yellow: karst areas (source: Chen et al. 2017). In orange: deserts (source: Olson & Dinerstein 2002). In violet, encircled by discontinued line: hypogean karst systems (source: Klimchouk 2007). 98. In blue: ice cover and permafrost extent during the Last Glacial Maximum (sources: Ehlers et al. 2011;Lindgren et al. 2016). 99. In blue: areas covered with either continuous or patched permafrost at present (source: Brown et al. 2002).
campodeids, three subfamilies are present in Asia. Lepidocampinae has twelve species in Asia and one cave-adapted species, L. hypogaea, known from seven caves in Sulawesi. Campodeinae has 52 soil species all across Asia, but only one genus Pacificampa is present in caves, three species in the Far East of Russia and northeast of China and two in Japan. It is Plusiocampinae, which has the lion's share of the Asian cave-dipluran diversity with eight genera and 12 species present in all Asian regions, but with only three species in soil habitats. Among cave Plusiocampinae, Plusiocampa and Vandelicampa occupy caves in western Asia in the Western Caucasus, Turkey and Lebanon; Turkmenocampa is found in a single cave of Central Asia in Turkmenistan; Simlacampa is known from southern Asia and was found in three caves of Penjab, India; Anisuracampa is known from a single cave in southeast Myanmar; whereas Hubeicampa gen. nov. and Whittencampa are found in eastern Asia, from Central to South China.
Asian cave-ecosystems have recently shown their remarkable biodiversity and extraordinary cave-adapted characteristics in many zoological groups, diplurans included (Deharveng et al. 2008;Golovatch 2015;Tian et al. 2016;Sendra et al. 2017aSendra et al. , 2018Sendra & Deharveng 2020), and the present contribution highlights the relevance of all cave-adapted diplurans in all regions from Asia.