Holotype ♂ (SCAU GD7), China, Guangdong Province, Qingyuan City, Yangshan County, Taiping Town, Niubi Village, Cave Niubi Yan, 24°10'23.93"N, 112°33'27.50"E, 100 m alt., 2014-XII-27, leg. Tian Mingyi, Liu Weixin, Huang Sunbin & Wang Xinhui. Paratypes: 1 ♂ (ZFMK), 1 ♂, 1 ♀ (SEM), 2 ♂, 1 ♀ (SCAU GD7), same data as the holotype.

To emphasise the apicomesal process of the anterior gonopod coxite being hook-shaped.

Differs from congeners of the formosus-group by the anterior gonopod showing a high and digitiform process, in which the tip is hook-shaped, coupled with a short, distally pectinate flagellum of the posterior gonopod. Based on molecular evidence, G. sinuatoprocessus Zhao & Liu, sp. nov. differs from all other Glyphiulus species analysed in a > 16.1% uncorrected p-distance of the COI barcoding gene.

Length ca. 33.0–42.0 (♂) or 45.0–50.0 mm (♀), mid-body rings round in cross-section, their width and height similar, 2.2–2.3 mm (♂) or 2.5–2.8 mm (♀). Body with 52–65 (♂) or 70–76 (♀) podous + 1–4 apodous rings + telson. Colouration orange-brownish to brownish, anterior part of body much darker in alcohol, red-brownish (Fig. 5A).

Figure 5. 

Pictures of live animals A G. sinuatoprocessus Zhao & Liu, sp. nov. from Cave Niubi Yan B G. scutatus Zhao & Liu, sp. nov. from Cave Bianfu Dong C G. portaliformis Zhao & Liu, sp. nov. from Cave Baiyan Dong D G. xiniudong Zhao & Liu, sp. nov. from Cave Xiniu Dong E G. conuliformis Zhao & Liu, sp. nov. from Cave Yanzi Dong.

Head surface smooth (Fig. 7A). Labrum with 4 or 5 teeth anteromedially. Ocellaria blackish, with 12–15 (♂) or 17–23 (♀) ommatidia arranged in 1–3 irregular linear rows (Fig. 7A). Antennae relatively long, reaching back to ring 3; in length, antennomeres 5 > 3 > 2 > 4 > 6 > 1 > 7. Antennomeres 5–7 each with a distodorsal field or corolla of bacilliform sensilla (sensory bacilli). Antennomere 7 with four sensory cones (Fig. 7A). Gnathochilarium with a separate promentum, lamellae linguales and promentum densely setose, a few setae on mentum (Fig. 7B). Mandible with a large external tooth and an internal tooth, the latter provided with nine cusps.

Collum : crests complete and evident; carinotaxy formula I–III+P+M (Fig. 7A). Following metaterga strongly crested; carinotaxy formula 2/2+I/i+3/3 (Fig. 6). Ozoporiferous tubercles very large, much higher than wide. Tegument delicately alveolate-areolate, fine longitudinal striations in front of stricture. Metatergal setae absent. Rings 2 and 3 with long pleural flaps (Fig. 6D). Limbus more or less regularly denticulate.

Figure 6. 

Glyphiulus sinuatoprocessus Zhao & Liu, sp. nov., ♂ paratype A rings 2–4, dorsal view B, E mid-body rings, dorsal and lateral views, respectively C posterior body rings, dorsal view D rings 2–5, sublateral view F cross-section of a mid-body ring, caudal view. Scale bars: 1 mm. Abbreviation: pf: pleural flap.

Figure 7. 

Glyphiulus sinuatoprocessus Zhao & Liu, sp. nov., ♂ paratype A head, collum and antenna B gnathochilarium C penes D leg 1, frontal view E leg 3, frontal view F mid-leg G ♀ paratype, vulvae H anterior gonopods, caudal view I posterior gonopod, caudal view J flagellum. Scale bars: 0.5 mm (A, B), 0.2 mm, (C–I), 0.1 mm (J).

Epiproct simple, very narrow, caudal edge uneven, with a strong central tubercle dorsally (Fig. 6C). Paraprocts rather regularly convex, each with several irregular rows of setae. Hypoproct transversely bean-shaped, with 3+3 small setae near caudal margin.

Legs slender, about 1.2 times as long as mid-body height; claw with a small accessory spine at base, about 1/3 as long as claw (Fig. 7F). ♂ legs 1 very strongly reduced, represented by a sternum showing a pair of small, fused, paramedian, subunciform prongs directed forward, with about 10–11+10–11 long setae at base; flanked by strongly separated, rudimentary, 2-segmented leg vestiges, first segment being much larger (Fig. 7D). ♂ legs 2 slightly hypertrophied, coxae large; penes small, much shorter than coxae, oblong-subtrapeziform, each with two strong setae distolaterally (Fig. 7C). ♂ legs 3 modified through coxae being especially slender and elongate (Fig. 7E). ♂ femora 6 and 7 normal, without modifications.

Anterior gonopods (Fig. 7H) with a broad and plate-shaped coxosternum supporting about 18–20 microsetae near distal margin; apicomesal process of coxite very high, digitiform, tip narrow and hook-shaped. Telopodite large and coiled, 1-segmented, lateral in position, slightly higher than lateral corner of coxite and with a field of 5–10 microspinules at base and 6–8 strong setae apically.

Posterior gonopods (Fig. 7I) compact. Coxite subquadrate, with a circular field of 12–15 basolateral microspinules in frontal view and an elongated field of 20–25 microspinules in caudal view; apical and mesal parts of coxite with dense, strong and curved setae. Lamelliform lobe membranous, with a short, distally pectinate flagellum (Fig. 7J).

Vulvae very simple, bare, modestly emarginate medially (Fig. 7G).

In the absence of direct troglomorphic traits, this species can only be considered as troglophilic at most. In the Guangdong Province and in Hong Kong, only G. formosus and G. granulatus have been recorded as yet.

Holotype ♂ (SCAU G24), China, Guangdong Province, Yangjiang City, Yangchun, Cave Yanzi Dong, 22°5'N, 111°36'50"E, 400 m alt., 2016-X-29, leg. Tian Mingyi, Chen Mengzhen & Wang Dianmei. Paratypes: 2 ♂, 26 ♀ (SCAU G24), same data as the holotype.

To emphasise the metatergal anterior tubercles being very sharp and coniform.

Differs from congeners of the formosus-group by the conical shape of the anterior tubercles of metaterga and by the first segment of the telopodite being significantly enlarged in ♂ legs 1, combined with the anterior gonopod process being slender, finger-shaped and curved inwards distally. Based on molecular evidence, G. conuliformis Zhao & Liu, sp. nov. differs from all other Glyphiulus species analysed in a > 17.0% uncorrected p-distance of the COI barcoding gene.

Length ca. 32.5–37.0 (♂) or 47.0–58.0 mm (♀), mid-body rings round in cross-section, their width and height similar, 1.7–2.0 mm (♂) or 2.2–3.0 mm (♀). Body with 48–53 (♂) or 57–67 (♀) podous + 2–4 apodous rings + telson. Colouration dark brownish, head and legs yellowish (Fig. 5E).

Head surface smooth. Labrum with four teeth anteromedially (Fig. 9C). Ocellaria blackish, with 12–22 ommatidia arranged in 2–3 irregular linear rows (Fig. 9A). Antennae slender, slightly clavate, reaching back to middle of rings 2 and 3; in length, antennomeres 5 > 3 > 2 ≈ 4 > 6 > 1 > 7. Antennomeres 5–7 each with a distodorsal field or corolla of bacilliform sensilla (sensory bacilli). Antennomere 7 with four sensory cones (Fig. 9B). Gnathochilarium with a separate promentum, polytrichous on promentum and mentum, lamellae linguales each with 6–7 setae (Fig. 9C). Mandible not dissected.

Collum : crests complete and evident; carinotaxy formula I–III+P+M (Figs 8A, B). Following metaterga strongly crested, anterior tubercles very sharp and coniform; carinotaxy formula 2/2+I/i+3/3 (Fig. 8). Ozoporiferous tubercles very large, coniform (Figs 8C, D). Tegument delicately alveolate-areolate, fine longitudinal striations in front of stricture. Rings 2 and 3 with long pleural flaps.

Figure 8. 

Glyphiulus conuliformis Zhao & Liu, sp. nov., ♂ paratype A, B anterior body rings, dorsal and lateral views C, D mid-body rings, dorsal and lateral views, respectively E, F posterior body rings, dorsal and ventral views, respectively. Scale bars: 1 mm. Abbreviation: ot: ozoporiferous tubercle.

Epiproct simple, caudal edge with a very low central protrusion, dorsally with an obvious sharp tubercle (Fig. 8E). Paraprocts regularly convex, each with several irregular rows of setae (Fig. 8F). Hypoproct transversely bean-shaped, with 3+3 small setae.

Legs slender, about 1.3 times as long as mid-body height; claw with a small accessory spine at base, about 1/6 as long as claw (Fig. 9D). ♂ legs 1 very strongly reduced, represented by a sternum showing a pair of small, unfused, paramedian, subunciform prongs directed forward, with about 4–5+4–5 long setae at base; flanked by strongly separated, rudimentary, 2-segmented, asymmetric leg vestiges, first segment significantly enlarged (Fig. 9F). ♂ legs 2 slightly hypertrophied, coxae large; penes rather small, much shorter than coxae, oblong-subtrapeziform, each with 1 or 2 strong setae distolaterally (Fig. 9H). ♂ legs 3 modified through coxae being especially slender and elongate (Fig. 9E). ♂ femora 6 and 7 normal, without modifications.

Figure 9. 

Glyphiulus conuliformis Zhao & Liu, sp. nov., ♂ paratype A head and collum B right antenna, oral view C gnathochilarium D mid-leg and claw E leg 3, caudal view F leg 1, frontal view G ♀ paratype, vulvae H penes I anterior gonopods, caudal view J, K posterior gonopods, frontal and caudal views, respectively. Scale bars: 0.5 mm (A–E), 0.2 mm (F–K).

Anterior gonopods (Fig. 9I) with a broad and plate-shaped coxosternum supplied with about 14–16 microsetae near distal margin; apicomesal process of coxite high, slender and digitiform, curved inwards distally. Telopodite very large and stout, coiled, 1-segmented, lateral in position, almost parallel to lateral corner of coxite, with a field of 8–16 microspinules at base and five strong setae apically.

Posterior gonopods (Figs 9J, K) compact. Coxite subtrapezoid, with a longitudinal field of 22–24 basolateral microspinules in frontal view and a slanted field of 32–36 median microspinules in caudal view; apical and mesal parts of coxite with dense, strong and curved setae. Lamelliform lobe membranous, with a rather short and broad flagellum.

Vulvae very simple, bare, M-shaped (Fig. 9G).

In the absence of direct troglomorphic traits, this species can only be considered as troglophilic at most.

Holotype ♂ (SCAU G78), China, Guangxi Zhuang Autonomous Region, Laibin City, Wuxuan County, Cave Xiniu Dong, 23°33'N, 109°32'55"E, 100 m alt., 2021-I-15, leg. Tian Mingyi, Liu Weixin & Zhao Yi. Paratypes: 4 ♂, 30 ♀ (SCAU G78), same data as the holotype.

To emphasise the provenance of this species from the “Xiniu” Cave, in Chinese meaning “rhinoceros”; noun in apposition.

Differs from congeners of the formosus-group by the metazonae with an obvious, corrugate, carved texture and by the leg claw with a rather large accessory spine, coupled with a long subtriangular coxosternum of the anterior gonopod, the latter process being narrow and digitiform. Based on molecular evidence, G. xiniudong Zhao & Liu, sp. nov. differs from all other Glyphiulus species analysed in a > 14.7% uncorrected p-distance of the COI barcoding gene.

Length ca. 28.0–31.0 (♂) or 26.0–41.0 mm (♀), mid-body rings round in cross-section, their width and height similar, 1.1–1.3 (♂) or 1.2–1.8 mm (♀). Body with 42–54 (♂) or 45–64 (♀) podous + 0–3 apodous rings + telson. Colouration purple-brownish, legs almost transparent (Fig. 5D).

Head surface smooth. Labrum with four teeth anteromedially. Ocellaria blackish, with 9–13 ommatidia arranged in 1–2 irregular linear rows (Figs 10B, 11A). Antennae short, slightly clavate, reaching back to caudal margin of ring 2; in length, antennomeres 5 > 3 > 4 ≈ 2 > 6 > 1 > 7. Antennomeres 5–7 each with a distodorsal field or corolla of bacilliform sensilla (sensory bacilli). Antennomere 7 with four sensory cones (Fig. 11B). Gnathochilarium with a separate promentum, polytrichous on promentum and mentum, lamellae linguales each with 6–7 setae (Fig. 11B). Mandible not dissected.

Figure 10. 

Glyphiulus xiniudong Zhao & Liu, sp. nov., ♂ paratype A, B anterior body rings, dorsal and lateral views C, D mid-body rings, dorsal and lateral views, respectively E, F posterior body rings, dorsal and ventral views, respectively. Scale bars: 1 mm.

Figure 11. 

Glyphiulus xiniudong Zhao & Liu, sp. nov., ♂ paratype A head and collum B gnathochilarium and left antenna C mid-body leg and claw D leg 1, frontal view E penes F ♀ paratype, vulvae G anterior gonopods, caudal view H, J posterior gonopod, frontal view I, K posterior gonopod, caudal view. Scale bars: 0.5 mm (A, B), 0.2 mm (C–I), 0.1 mm (J, K).

Collum : crests complete and evident; carinotaxy formula I–III+P+M (Figs 10A, 11A). Following metaterga strongly crested; carinotaxy formula 2/2+I/i+3/3 (Fig. 10). Ozoporiferous tubercles very large, coniform. Prozonae delicately alveolate-areolate, fine longitudinal striations in front of stricture. Metazonae with an obvious, corrugate, carved texture (Fig. 10). Rings 2 and 3 with long pleural flaps.

Epiproct simple, caudal edge uneven, with an obvious central tubercle dorsally (Fig. 10E). Paraprocts regularly convex, each with several irregular rows of setae. Hypoproct transversely bean-shaped, with 4+4 small setae (Fig. 10F).

Legs short, about as long as mid-body height; claw with a relatively large accessory spine at base, about half as long as claw (Fig. 11C). ♂ legs 1 very strongly reduced, represented by a sternum showing a pair of small, more slender, fused, paramedian, subunciform prongs directed forward, with about 7–9+7–9 long setae at base; flanked by strongly separated, rudimentary, 1-segmented leg vestiges, with some setae (Fig. 11D). ♂ legs 2 slightly hypertrophied, coxae large; penes rather small, much shorter than coxae, oblong-subtrapeziform, each with three strong setae distolaterally (Fig. 11E). ♂ legs 3 modified through coxae being especially slender and elongate. ♂ femora 6 and 7 normal, neither modifications.

Anterior gonopods (Fig. 11G) with a long subtriangular coxosternum with about 9–11 microsetae near distal margin; apicomesal process of coxite very high, rather narrow and digitiform, modestly curved inwards. Telopodite very large, stout and coiled, 1-segmented, lateral in position, much higher than lateral corner of coxite, with a field of six microspinules at base and 6–7 strong setae apically.

Posterior gonopods (Figs 11H, I) compact. Coxite subquadrate, with a circular field of about 10–15 basolateral microspinules in frontal view; with a field of 19 median microspinules and an apicolateral very strong and long seta in caudal view; apical and mesal parts of coxite with dense, strong and curved setae. Lamelliform lobe membranous, with a rather small, spine-like flagellum.

Vulvae very simple, bare, modestly emarginate medially (Fig. 11F).

In the absence of direct troglomorphic traits, this species can only be considered as troglophilic at most.

Holotype ♂ (SCAU WL30), China, Guangxi Zhuang Autonomous Region, Hechi City, Du’an Yao Autonomous County, Napang Dong, 24°08'22"N, 107°51'07"E, 650 m alt., 2015-VII-26, leg. Chen Jujian, Wang Xinhui & Tang Mingruo. Paratypes: 2 ♂, 12 ♀ (SCAU WL30), same data as the holotype. 3 ♂, 4 ♀ (SCAU G72), same County, Cave Bianfu Dong, 24°01'55"N, 108°20'12"E, 550 m alt., 2017-VIII-18, leg. Tian Mingyi, Huang Sunbin, Wang Dianmei & Chen Mengzhen.

To emphasise the anterior gonopod showing a scalloped and shield-shaped coxosternum.

Differs from congeners of the formosus-group by both ♂ femora 6 and 7 being slightly inflated and the anterior gonopod without apicomesal process, as well as the posterior gonopod with less than ten microspinules. Based on molecular evidence, G. scutatus Zhao & Liu, sp. nov. differs from all other Glyphiulus species analysed in a > 15.8% uncorrected p-distance of the COI barcoding gene.

Length of both sexes ca. 25.0–41.0 mm, mid-body rings round in cross-section, their width and height similar, 1.3–1.8 (♂) or 1.8–2.0 mm (♀). Body with 55–68 podous + 1–4 apodous rings + telson. Colouration purple-brownish to dark brownish, legs light brownish to almost transparent (Fig. 5B).

Head surface smooth. Labrum with four teeth anteromedially (Fig. 13A). Ocellaria blackish, with 8–12 ommatidia arranged in two irregular linear rows (Fig. 12B). Antennae short, slightly clavate, reaching back to ring 3; in length, antennomeres 5 > 3 > 4 > 2 > 6 > 1 > 7. Antennomeres 5–7 each with a distodorsal field or corolla of bacilliform sensilla (sensory bacilli). Antennomere 7 with four sensory cones (Fig. 13B). Gnathochilarium with a separate promentum, setae variable in number, polytrichous or smooth on promentum and mentum, lamellae linguales each with 4–6 setae (Fig. 13A). Mandible not dissected.

Figure 12. 

Glyphiulus scutatus Zhao & Liu, sp. nov., ♂ paratype from Cave Napang Dong A, B anterior body rings, dorsal and lateral views C, D mid-body rings, dorsal and lateral views, respectively E, F posterior body rings, dorsal and ventral views, respectively. Scale bars: 1 mm.

Figure 13. 

Glyphiulus scutatus Zhao & Liu, sp. nov., ♂ paratype from Cave Napang Dong A gnathochilarium and right antenna B antenna tip C leg 3, frontal view D leg 1, frontal view E ♀ paratype, vulvae F penes G, J posterior gonopod, frontal and caudal views, respectively H mid-body leg and claw I anterior gonopods, caudal view. Scale bars: 0.5 mm (A, H), 0.2 mm (B–E), 0.1 mm (F, G, I, J).

Collum : crests complete and evident; carinotaxy formula I–III+P+M (Figs 11A, B). Following metaterga strongly crested and extremely sharp; carinotaxy formula 2/2+I/i+3/3 (Fig. 12). Ozoporiferous tubercles very large, subcylindrical. Prozonae delicately alveolate-areolate, fine longitudinal striations in front of stricture. Metazonae with an obvious, corrugate, carved texture (Fig. 12). Rings 2 and 3 with long pleural flaps.

Epiproct simple, very narrow, caudal edge uneven, with a clear central tubercle dorsally (Fig. 12E). Paraprocts regularly convex, each with several irregular rows of setae. Hypoproct transversely bean-shaped, with 3+3 small setae (Fig. 12F).

Legs slender, about 1.1–1.3 times as long as mid-body height; claw with a small accessory spine at base, about ¼ as long as claw (Fig. 13H). ♂ legs 1 very strongly reduced, represented by a sternum showing a pair of small, fused, paramedian, subunciform prongs directed forward, with 3+3 long setae at base; flanked by strongly separated, rudimentary, 2-segmented leg vestiges, with either a few or no setae (Fig. 13D). ♂ legs 2 slightly hypertrophied, coxae large; penes small, much shorter than coxae, oblong-subtrapeziform, each with 1–4 strong setae distolaterally (Fig. 13F). ♂ legs 3 modified through coxae being especially slender and elongate (Fig. 13C). Both ♂ femora 6 and 7 slightly inflated.

Anterior gonopods (Fig. 13I) with a scalloped and shield-shaped coxosternum with about 6–8 microsetae near lateral corner of coxite. Telopodite very large and coiled, 1-segmented, lateral in position, much higher than lateral corner of coxite, with a field of 5–7 microspinules at base and 4–5 strong setae apically.

Posterior gonopods (Figs 13G, J) compact. Coxite subquadrate, with a field of about 8–10 basolateral microspinules in frontal view and with a field of ten median microspinules in caudal view; apical and mesal parts of coxite with dense, strong and curved setae. Lamelliform lobe membranous, with a short, broad, distally spike-like flagellum.

Vulvae very simple, bare, rather faintly emarginate medially (Fig. 13E).

In the absence of direct troglomorphic traits, this species can only be considered as troglophilic at most.

Holotype ♂ (SCAU WL40), China, Guangxi Zhuang Autonomous Region, Hechi City, Bama County, Cave Baiyan Dong, 24°03'40"N, 107°08'16"E, 400 m alt., 2015-VII-31, leg. Chen Jujian, Wang Xinhui & Tang Mingruo. Paratypes: 7 ♂, 18 ♀ (SCAU WL40), same data as the holotype.

To emphasise the coxosternum of the anterior gonopods being portal-shaped.

Differs from congeners of the formosus-group by the epiproct showing a small caudal protrusion and the anterior gonopods being portal-shaped, combined with a foliate flagellum of the posterior gonopod. Based on molecular evidence, G. portaliformis Zhao & Liu, sp. nov. Differs from all other Glyphiulus species analysed in a > 14.4% uncorrected p-distance of the COI barcoding gene.

Length of both sexes ca. 41.0–56.0 mm, mid-body rings round in cross-section, their width and height similar, 2.2–3.0 mm. Body with 55–67 podous + 1 apodous ring + telson. Colouration brownish, legs almost transparent (Fig. 5C).

Head surface smooth. Labrum with 4 teeth anteromedially (Fig. 15A). Ocellaria blackish, with 14–17 ommatidia arranged in 2–3 irregular linear rows (Figs 14B, 15B). Antennae short, slightly clavate, reaching back to ring 4; in length, antennomeres 5 > 3 > 2 > 4 > 6 > 1 > 7. Antennomeres 5–7 each with a distodorsal field or corolla of bacilliform sensilla (sensory bacilli). Antennomere 7 with four sensory cones (Fig. 15A). Gnathochilarium with a separate promentum, polytrichous on promentum and mentum, lamellae linguales each with 6–7 setae (Fig. 15A). Mandible not dissected.

Figure 14. 

Glyphiulus portaliformis Zhao & Liu, sp. nov., ♂ paratype A, B anterior body rings, dorsal and lateral views C, D mid-body rings, dorsal and lateral views, respectively E, F posterior body rings, dorsal and ventral views, respectively. Scale bars: 1 mm.

Figure 15. 

Glyphiulus portaliformis Zhao & Liu, sp. nov., ♂ paratype A gnathochilarium and left antenna B head and collum C penes D mid-leg and claw E leg 3, caudal view F leg 1, frontal view G ♀ paratype, vulvae H anterior gonopods, caudal view I, J posterior gonopod, caudal and frontal views, respectively. Scale bars: 0.5 mm (A, B, D, E), 0.2 mm (C, F–J).

Collum : crests complete and evident; carinotaxy formula I–III+P+M (Figs 14A, B, 15B). Following metaterga strongly crested; carinotaxy formula 2/2+I/i+3/3 (Fig. 14). Ozoporiferous tubercles very large, coniform. Prozonae delicately alveolate-areolate, fine longitudinal striations in front of stricture. Metazonae with an obvious, corrugate, carved texture (Fig. 14). Rings 2 and 3 with long pleural flaps.

Epiproct simple, caudal edge with a central conical protrusion and dorsally with a strong central rounded tubercle (Fig. 14E). Paraprocts regularly convex, each with several irregular rows of setae. Hypoproct transversely bean-shaped, no setae visible (Fig. 14F).

Legs slender, about 1.1 times as long as mid-body height; claw with a small accessory spine at base, about 1/5 as long as claw (Fig. 15D). ♂ legs 1 very strongly reduced, represented by a sternum showing a pair of small, fused, paramedian, subunciform prongs directed forward, with about ten long setae at base; flanked by strongly separated, rudimentary, 2-segmented leg vestiges, second segment very large, stout, rod-shaped (Fig. 15F). ♂ legs 2 slightly hypertrophied, coxae large; penes small, much shorter than coxae, oblong-subtrapeziform, each with two or three strong setae distolaterally (Fig. 15C). ♂ legs 3 modified through coxae being especially slender and elongate (Fig. 15E). Both ♂ femora 6 and 7 normal, without modifications.

Anterior gonopods (Fig. 15H) very tall, with a portal-shaped coxosternum with about 16–18 microsetae near distal margin; apicomesal process of coxite subtriangular, tip small, slightly curved inwards. Telopodite very large and clavate, 1-segmented, lateral in position, slightly higher than lateral corner of coxite, with a field of nine microspinules at base and six strong setae apically.

Posterior gonopods (Figs 15I, J) compact. Coxite nearly parallelogram-shaped, with an elongated field of about 14 basolateral microspinules in frontal view and with a field of 16 median microspinules in caudal view; apical and mesal parts of coxite with dense, strong and curved setae. Lamelliform lobe membranous, with a relatively long, broad, distally foliate flagellum.

Vulvae very simple, bare, faintly emarginate medially (Fig. 15G).

In the absence of direct troglomorphic traits, this species can only be considered as troglophilic at most.

4 ♂, 7 ♀ (SCAU WL37), China, Guangxi Zhuang Autonomous Region, Hechi City, Fengshan County, Zhaiya Town, Cave Jianbang Dong, 24°43'02.96"N, 107°13'11.21"E, 350 m alt., 2015-VIII-4, leg. Chen Jujian, Huang Sunbin & Tang Mingruo.

This species has been described from the Cave Xianren Dong, Bala Town, Tian’e County, Guangxi, China. The new samples were collected from a cave in the neighbouring county, both being located close geographically (Fig. 16). The above material is in good agreement with the original description by Jiang et al. (2018), while intraspecific p-distance is 2.1%, based on DNA-barcoding. Based on molecular evidence, G. impletus differs from all other Glyphiulus species analysed from between 11.9% (compared to G. calceus) and 23.6% (compared to G. duangdee).

Figure 16. 

A distribution map of the Glyphiulus formosus species group in China. 1 G. recticullus 2 G. foetidus 3 G. impletus 4 G. calceus 5 G. fortis 6 G. portaliformis Zhao & Liu, sp. nov. 7 G. pulcher 8 G. scutatus Zhao & Liu, sp. nov. 9 G. echinoides 10 G. xiniudong Zhao & Liu, sp. nov. 11 G. sinuatoprocessus Zhao & Liu, sp. nov. 12 G. conuliformis Zhao & Liu, sp. nov. 13 G. formosus 14 G. hainanensis. Abbreviations: FJ: Fujian, HN: Hainan, HuN: Hunan, GD: Guangdong, GX: Guangxi, GZ: Guizhou, ZJ: Zhejiang.

9 ♂ (SCAU WL38), China, Guangxi Zhuang Autonomous Region, Hechi City, Fengshan County, Jiangzhou Underground Corridor, 24°30'4"N, 106°53'46"E, 850 m alt., 2013-VI-30, leg. Tian Mingyi, Liu Weixin, Lin Wei, Yin Haomin & Huang Sunbin. 4 ♂, 2 ♀ (SCAU WL39), China, Guangxi Zhuang Autonomous Region, Baise City, Linyun County, Cave Fengniu Dong, 24°28'39"N, 106°37'52"E, 700 m alt., 2017-VI-9, leg. Tian Mingyi, Liu Weixin, Wang Xinhui & Tang Mingruo.

This species has been found to have a relatively wide distribution in Guangxi, involving the Lingyun, Fengshan, Nandan and Donglan Counties. The new samples add only the records of two cave collections (Fig. 16). The above material is in good agreement with the original description by Jiang et al. (2018), while intraspecific p-distances are 2.4–8.2%, based on DNA-barcoding. Based on molecular evidence, G. impletus differs from all other Glyphiulus species analysed from between 11.9% (compared to G. calceus) and 24.9% (compared to G. foetidus).