Durgella pentata, Pholyotha & Sutcharit & Lwin & Panha, 2024

Pholyotha, Arthit, Sutcharit, Chirasak, Lwin, Ngwe & Panha, Somsak, 2024, Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population., Zoological Studies 63 (14), pp. 141-149 : 13-26

Durgella rhaphiellus ( Martens, 1867) ( Fig. 8A )

Helicarion rhaphiellus Martens, 1867: 69 , pl. 12, fig. 9. Type locality: Siam [ Thailand]. Tryon 1885: 177, pl. 40, figs. 36–38. Fischer and Dautzenberg 1904: 392. Saurin 1953: 113.

Durgella rhaphiellus – Inkhavilay et al. 2019: 83, fig. 39c.

Material examined: Syntypes ZMB / MOLL 5033 (2 shells; Fig. 8A ) from Siam [ Thailand].

Diagnosis: Shell subglobose, fragile, polished, pale golden-amber, and large body whorl very descending in front.

Description: Shell ( Fig. 8A ). Shell subglobose, small (width 7.5 mm, height 5 mm), thin, smooth, polished, pale golden-amber. Spire slightly elevated; suture slightly impressed. Whorls 3, convex and rapidly increasing; body whorl enlarged, well-rounded and descending. Aperture oblique, roundly lunate, width greater than height; peristome thin. Columellar margin simple and slightly expanded near umbilicus. Umbilicus minute.

Distribution: Durgella rhaphiellus is only known from the type locality in Thailand, and its exact locality is unknown.

Remarks: This species was described by Martens (1867) based on two specimens with ‘Siam’ as the type locality, which is very rough and covers a wide geographical area. However, the itinerary of E. von Martens indicates that he arrived in mainland Southeast Asia and made stops in ‘Bangkok’ and ‘Petshaburi’ for collecting specimens over a period of around three months ( Martens 1876; Kabat and Boss 1997). This suggests that ‘Petshaburi’ [= Phetchaburi Province, Thailand] is the collection locality of this species, which is much more precise than ‘Siam’. Since this species was first discovered, there have been no additional specimens collected, records, or literature references. Unfortunately, in our surveys we could not find any specimens that could be identified as this species for redescription.

Without further evidence of the soft-part anatomy depicted from the topotype; the taxonomic status and relationship of D. rhaphiellus is still provisional; therefore, we recognise it as valid species per Inkhavilay et al. (2019). However, D. rhaphiellus differs from D. levicula from peninsular Thailand by having a relatively smaller size and more descending last whorl. In addition, the apertural width is greater than the height in D. rhaphiellus , whereas these parameters are more equal in D. levicula .

Durgella siamensis Möllendorff, 1902 ( Figs. 1 , 3E , 8B–D , 9G, H , 10B, C , 11G, H )

Durgella siamensis Möllendorff, 1902: 155 . Type locality: Muoklek, Siam [Muak Lek District, Saraburi Province, Thailand]. Hemmen and Hemmen 2001: 43.

Bensonia (Durgella) siamensis – Fischer and Dautzenberg 1904: 395.

Material examined: Syntype SMF 227168/1 (1 shell) from Muoklek, Siam [Muak Lek District, Saraburi Province, Thailand]. THAILAND: Sarika Waterfall, Mueang District, Nakhon Nayok Province (14°18'31.3"N, 101°15'17.9"E): CUMZ 14232 (10 preserved specimens).

Diagnosis: Shell subglobose to globose, fragile, polished, dark yellow with a creamy tinge, and large body whorl very descending in front; animal with blackish body and four mantle extensions; genitalia with very short vagina, rather long epiphallus, short epiphallic caecum, moderate gametolytic duct, and rather large dart apparatus.

Description: Shell ( Figs. 8B–D , 9G, H ). Shell subglobose to globose, small (width up to 9.3 mm, height up to 6.5 mm), transparent, smooth, thin, polished, and yellowish-corneous or dark yellow with creamy tinge. Protoconch smooth ( Fig. 9G, H ). Spire slightly elevated; suture slightly impressed. Whorls 3½– 4, convex, rapidly increasing, last whorl enlarged, well-rounded, and descending. Aperture oblique, diagonal, roundly lunate, width greater than height; peristome thin. Columellar margin thin and little expanded near umbilicus. Umbilicus minute.

Genital organs ( Fig. 10B, C ). Atrium (at) enlarged and very short. Penis (p) long, cylindrical, and with thin penial sheath encircling from atrium to almost middle of penis. Inner sculpture of penis uniform throughout with small conical penial pilasters (pp). Epiphallus (e1 + e2) cylindrical and slightly longer than penis; inner sculpture of e1 and e2 with thin longitudinal folds, and one thickened longitudinal pilaster. Epiphallic caecum (ec) short and small; penial retractor muscle (prm) thick and attached at tip of epiphallic caecum. Vas deferens (vd) long and thin tube. Vagina (v) very short or inconspicuous. Dart apparatus (da) short, enlarged cylindrical, and connected to atrium chamber. Gametolytic duct (gd) slightly shorter than penis, proximally enlarged at base and then tapering distally; gametolytic sac (gs) enlarged and bulbous. Free oviduct (fo) slightly shorter than penis; proximal part enlarged and bulbous; distal end large and cylindrical. Oviduct enlarged lobules; prostate gland running alongside oviduct.

Radula ( Fig. 11G, H ). Teeth pectiniform, arrangement and shape similar to those of D. levicula , and with half row ca. 380 teeth. Central tooth very reduced to tiny unicuspid. Lateromarginal teeth elongated bicuspid, outer edge serrated with 3 to 6 cusps. Outermost teeth shorter than inner teeth.

External appearance ( Fig. 3E ). Animal with fleshy-grey on anterior body; posterior body blackish; foot margin pale grey. Four mantle extensions well-developed and dark grey, graduating to cream near foot margin. Caudal horn raised, large, and blackish.

Distribution: The type locality and the newly collected sample suggest that D. siamensis is probably restricted to the Saraburi and Nakhon Nayok provinces, central Thailand ( Fig. 1 ).

Remarks: Durgella siamensis was originally described by O.F. von Möllendorff (1902) based on specimens collected by H. Fruhstorfer, a butterfly collector, from Muoklek [Muak Lek District, Saraburi Province], central Thailand. In this study, we collected specimens from Nakhon Nayok Province, very close to the type locality, and found that D. siamensis tended to vary in shell shape between the subadult ( Fig. 8C ) and fully grown ( Fig. 8D ) specimens. The subadult shell of this species is similar to D. rhaphiellus ( Fig. 8A ), but the latter species differs by having a more depressed and slightly smaller shell. Because of the shell similarity between D. siamensis and D. rhaphiellus , an anatomical investigation and molecular study are necessary to clarify their taxonomic status.

Durgella siamensis differs from D. levicula , a common and widespread species in Thailand, by the descending last whorl. Anatomically, D. siamensis has thin longitudinal folds on the inner sculpture of the epiphallus (e2) ( Fig. 10C ), whereas D. levicula has small longitudinal dash-like pilasters varying in length ( Fig. 6C ) on the inner sculpture of the epiphallus (e2).

Durgella libas Solem, 1966 ( Figs. 1 , 3F , 4A , 12A, B , 13A, B , 14 , 17A, B )

Durgella libas Solem, 1966: 50 , figs. 7–9, 13b. Type locality: Wang Dao, North Thailand [Chiang Dao District, Chiang Mai Province, Thailand]. Panha 1996: 35. Hemmen and Hemmen 2001: 43. Inkhavilay et al. 2019: 83, figs. 19f, 57c.

Material examined: THAILAND: Wachirathan Waterfall, Chom Thong District, Chiang Mai Province (18°32'30.7"N, 98°35'53.6"E): CUMZ 14236 (4 preserved specimens). Phra-Ruesi Cave, Mueang District, Chiang Mai Province (18°48'24.5"N, 98°54'38.3"E): CUMZ 14237 (4 preserved specimens). San Ku, Mueang District, Chiang Mai Province (18 ° 48 ' 56.8 " N, 98 ° 53 ' 38.7 "E): CUMZ 14238 (7 preserved specimens). Doi Suthep-Pui Nature Trail, Mueang District, Chiang Mai Province (18°49'23.4"N, 98°53'23.6"E): CUMZ 14235 (1 preserved specimen).

Diagnosis: Shell depressedly subglobose, membranous, polished, dark yellow with a creamy tinge; animal with four mantle extensions; genitalia with slightly long and enlarged penis, rather long epiphallus, small epiphallic caecum, short gametolytic duct, and very large dart apparatus.

Description: Shell ( Figs. 12A, B , 13A, B ). Shell depressedly subglobose, small to medium (width up to 13.0 mm, height up to 7.6 mm), transparent, very thin (membranous) and fragile, polished, and dark yellow with creamy tinge. Protoconch surface sculptured with faint spiral furrows with regular malleation ( Fig. 13A, B ). Spire slightly elevated; suture slightly impressed. Whorls 4–4½, convex and rapidly increasing; last whorl smooth, enlarged, and well-rounded. Aperture oblique, diagonal, roundly lunate, broader than high, and apertural lip simple. Columellar margin thin and little expanded near umbilicus. Umbilicus minute.

Genital organs ( Fig. 14 ). Atrium (at) enlarged and very short. Penis (p) slightly long, enlarged with irregular shape, and with thin penial sheath encircling from atrium to almost middle of penis. Inner sculpture of penis uniform throughout with small conical penial pilasters (pp). Epiphallus (e1 + e2) same as penis length but its diameter much narrower; e1 rather long and slender; e2 shorter and bulbous. Inner sculpture: e1 with small longitudinal dash-like pilasters, and e2 with one large longitudinal pilaster. Epiphallic caecum (ec) short, small, and oblong; penial retractor muscle (prm) thick and attached at tip of epiphallic caecum. Vas deferens (vd) long and thin tube. Vagina (v) very short or nearly absent. Dart apparatus (da) rather short, enlarged cylindrical, and connected to atrium chamber. Gametolytic duct (gd) proximally enlarged and tapering to smaller tube distally; gametolytic sac (gs) enlarged and bulbous. Free oviduct (fo) slightly shorter than penis; proximally enlarged and bulbous; distal part cylindrical. Oviduct enlarged lobules; prostate gland running alongside oviduct.

Radula ( Fig. 17 A, B ). Teeth pectiniform, arrangement and shape similar to those of D. levicula , and with half row ca. 430 teeth. Central tooth very reduced to tiny unicuspid. Lateromarginal teeth elongated bicuspid and outer edge serrated with several tiny cusps. Outermost teeth shorter than inner teeth.

External appearance ( Figs. 3F , 4A ). Posterior body with pale reddish-brown to light yellow-brown on dorsal and light yellow near foot margin; anterior body light yellow or creamy. Four mantle extensions well-developed and creamy. Caudal horn raised, large, and reddish-brown.

Distribution: The species is known from Chiang Mai Province, northern Thailand ( Fig. 1 ) and the newly collected samples suggest that this species prefers habitats in high mountainous areas.

Remarks: Solem (1966) listed the specimens from Ban Kao and Sai Yok [both from Kanchanaburi Province, western Thailand] as this species. However, we have collected many Durgella specimens from these two localities and found that the soft body colour, shells, and genital morphology are as those of D. levicula instead. Therefore, we conclude that D. libas is probably restricted to northern Thailand.

Durgella pentata Pholyotha & Panha , sp. nov. ( Figs. 1 , 3G , 4B , 12C, D , 13C, D , 15 , 17C, D ) urn:lsid:zoobank.org:act:398E44AA-406A-4C4C-9BE8-4BC191484BBF

Type material: Holotype CUMZ 14239 ( Fig. 12C ; width 11.5 mm, height 6.7 mm) . Paratypes same locality as holotype: CUMZ 14240 (52 preserved specimens; Fig. 12D ; width 11.0 mm, height 6.5 mm), 14241 (24 preserved specimens), NHMUK (2 shells).

Other material examined: THAILAND: Wat Tham Klaeb (Wat Tham Nguang Chang), Chiang Dao District, Chiang Mai Province (19°33'32.3"N, 99°03'46.2"E): CUMZ 14242 (7 preserved specimens). Pha Daeng Cave-Luang Cave, Chiang Dao District, Chiang Mai Province (19°34'40.7"N, 99°03'43.4"E): CUMZ 14243 (1 preserved specimen). Ban Pang Mayao, Chiang Dao District, Chiang Mai Province (19°26'10.5"N, 99°04'09.2"E): CUMZ 14244 (20 preserved specimens).

Type locality: Tham Chiang Dao, Chiang Dao District, Chiang Mai Province, Thailand (19°23'37.5"N, 98°55'39.6"E).

Diagnosis: Shell depressedly subglobose, thin, polished, dark yellow with a creamy tinge, and aperture roundly lunate, broader than high; animal with five mantle extensions; genitalia with very short vagina, small epiphallic caecum, short gametolytic duct, and very large dart apparatus.

Description: Shell ( Figs. 12C, D , 13C, D ). Shell depressedly subglobose, small to medium (width up to 11.7 mm, height up to 7.2 mm), transparent, polished, smooth, and yellowish-corneous or dark yellow with a creamy tinge. Protoconch surface sculptured with faint spiral furrows and regular strong malleation ( Fig. 13C, D ). Spire slightly elevated; suture impressed. Whorls 4–4½, convex, rapidly increasing; last whorl enlarged and well-rounded. Aperture oblique, diagonal, roundly lunate, width greater than height; peristome simple. Columellar edge thin and little expanded near umbilicus. Umbilicus minute.

Genital organs ( Fig. 15 ). Atrium (at) enlarged and very short. Penis (p) long, cylindrical, and with thick penial sheath encircling about half of penis length. Inner sculpture of penis with longitudinal penial ridges from proximal to near middle and then transforming to small conical penial pilasters (pp). Epiphallus (e1 + e2) slightly shorter than penis; e1 cylindrical and narrower than penis; e2 short and bulbous shape. Inner sculpture: e1 with one thickened longitudinal fold running from proximal to near middle; e2 distally with dense papillae, and one thickened longitudinal pilaster. Epiphallic caecum (ec) short and small; penial retractor muscle (prm) thick and attached at tip of epiphallic caecum. Vas deferens (vd) long and slender tube. Vagina (v) very short or inconspicuous. Dart apparatus (da) very large, long cylindrical, and connected to atrium chamber. Gametolytic duct (gd) short, about half of penis length, proximally enlarged and then tapering to smaller and slender tube; gametolytic sac (gs) enlarged and bulbous. Free oviduct (fo) shorter than penis; proximally enlarged and bulbous; distal part shorter and cylindrical. Oviduct enlarged lobules; prostate gland running alongside oviduct.

Radula ( Fig. 17C, D ). Teeth pectiniform and numerous with half row ca. 380 teeth. Central tooth reduced to bicuspid. Lateromarginal teeth elongated bicuspid, unclear separation between endocone and ectocone and outer edge serrated with inconspicuous cusps. Outermost marginal teeth shorter, smaller, and more conspicuous serrated cusps than inner teeth.

External appearance ( Figs. 3G , 4B ). Dorsal of posterior body yellow to dark brown and then transforming to light yellow below foot margin; anterior body light yellowish. Five mantle extensions well-developed and creamy. Caudal horn raised, large, and yellowish-brown to dark brown.

Etymology: The specific name “ pentata ” is from the Greek word meaning five, referring to the presence of five mantle extensions.

Distribution: The new species is known from Tham Chiang Dao and the vicinity of Chiang Dao District, Chiang Mai Province, northern Thailand ( Fig. 1 ).

Remarks: Durgella pentata sp. nov. differs from D. levicula , D. erratica , D. siamensis , and D. libas in the presence of five mantle extensions, while the other species have four mantle extensions. This new species also differs from the others by having a protoconch sculptured with strong malleation, while D. levicula , D. erratica , and D. siamensis have rather smooth or faint spiral furrows without malleation, and D. libas has spiral furrows with faint malleation.

Compared with D. libas , the species with the nearest geographic distribution, this new species possesses a smaller penis, shorter epiphallus, and inner penile structure with proximal longitudinal ridges transforming to distal small conical shapes. In contrast, D. libas has a larger penis, longer epiphallus, and inner penile sculpture with small conical shapes throughout the entire chamber.

In contrast to the other species with five mantle lobes ( D. birmanica and D. concinna ), this new species has a larger shell, lower spire, and more enlarged dart apparatus than D. birmanica . Compared with D. concinna , this new species has a larger and thinner shell with a minute umbilicus, and without a descending body whorl near the aperture.

Durgella nulla Pholyotha & Panha , sp. nov. ( Figs. 1 , 3H , 4C, D , 12E, F , 13E, F , 16 , 17E , F) urn:lsid:zoobank.org:act:0C470A9B-2B16-489F-B8BA-017009FACC56

Type material: Holotype CUMZ 14227 ( Fig. 11E ; width 12.9 mm, height 7.6 mm) . Paratypes same locality as holotype: CUMZ 14228 (19 preserved specimens; Fig. 11F ; width 12.9 mm, height 7.1 mm), NHMUK (2 shells).

Other material examined: THAILAND: Wat Tham Pha Pu, Mueang District, Loei Province (17 °34 '43.4 "N, 101 °42' 37.2 "E): CUMZ 14229 (8 preserved specimens). Wat Tham Pha Sing, Wang Saphung District, Loei Province (17°17'28.0"N, 101°49'40.1"E): CUMZ 14230 (1 preserved specimen). Wat Tham Phupha Thamsathit, Suwannakhuha District, Nong Bua Lam Phu Province (17°37'38.4"N, 102 °10 ' 15.3 " E): CUMZ 14231 (12 preserved specimens).

Type locality: Phu Pha Lom, Mueang District, Loei Province, Thailand (17°33'15.9"N, 101°52'04.9"E).

Diagnosis: Shell depressedly subglobose, membranous, polished, dark yellowish with olive tinge; animal with two different colours: upper body dark brown or blackish and from lower body to foot margin creamy; four mantle extensions; genitalia with very large penis, short gametolytic duct, and without dart apparatus.

Description: Shell ( Figs. 12E, F , 13E, F ). Shell depressedly subglobose, medium to large (width up to 13.5 mm, height up to 7.6 mm), transparent, smooth, very thin and fragile, polished, and dark yellowish with olive tinge. Protoconch surface sculptured with strong spiral furrows with or without malleation on furrows ( Fig. 13E, F ). Spire slightly elevated; suture little impressed. Whorls 4¼–4½, convex, rapidly increasing; body whorl enlarged and rounded; aperture oblique, roundly lunate, diagonal, width greater than height; peristome simple. Columellar margin thin and little expanded near umbilicus. Umbilicus minute.

Genital organs ( Fig. 16 ). Atrium (at) enlarged and very short. Penis (p) long, enlarged cylindrical, and with thin penial sheath encircling about half of penis length. Inner sculpture of penis with uniformly small conical penial pilasters (pp) throughout. Epiphallus (e1 + e2) shorter than half of penis length; e1 more slender and narrower than penis; e2 shorter and bulbous in shape. Inner sculpture: e1 with small conical knobs arranged on longitudinal folds, and one very large and smooth longitudinal fold; e2 distally with dense papillae, and one large longitudinal fold. Epiphallic caecum (ec) short, slightly enlarged; penial retractor muscle (prm) thickened and attached at tip of epiphallic caecum. Vas deferens (vd) long and thin tube. Vagina (v) long and enlarged cylindrical. Dart apparatus (da) absent. Gametolytic duct (gd) short, proximally enlarged and then tapering to a small tube; gametolytic sac (gs) enlarged and bulbous shape. Free oviduct (fo) about half of penis length, proximally enlarged, and distally cylindrical. Oviduct enlarged lobules; prostate gland running alongside oviduct.

Radula ( Fig. 17E, F ). Teeth pectiniform and numerous with half row ca. 460 teeth. Central tooth very reduced to tiny bicuspid. Lateromarginal teeth elongated bicuspid and outer edge serrated with 3 to 6 tiny cusps, gradually diminishing in size outwards. Outermost marginal teeth shorter, smaller, and with more conspicuous serrated cusps than inner teeth.

External appearance ( Figs. 3H , 4C, D ). Posterior body with dark brown or blackish on dorsal and immediately transforming to yellow or cream near foot margin; anterior body light yellowish-brown to dark brown. Four mantle extensions well-developed and light cream in colour. Caudal horn raised, large, and blackish.

Etymology: The specific name “ nulla ” is from the Latin adjective meaning not any, none, nobody, or no, and refers to the absence of dart apparatus.

Distribution: The new species is known from Loei and Nong Bua Lam Phu provinces in northeastern Thailand ( Fig. 1 ).

Remarks: Comparing the two new species described herein, D. nulla sp. nov. has four mantle extensions and no dart apparatus, whereas D. pentata sp. nov. has five mantle extensions and a large dart apparatus. Durgella nulla sp. nov. also differs from all other species from Thailand and Myanmar in the absence of a dart apparatus, whereas this organ appears in all other recognised species examined in this study.

Among species without a dart apparatus, D. assamica from Assam and D. rogersi from the Andaman Islands have a smaller subglobose shell with a long gametolytic organ ( Godwin-Austen 1881; Blanford and Godwin-Austen 1908), whereas D. nulla sp. nov. possesses a larger and depressedly subglobose shell with a short gametolytic organ and is locally endemic to Thailand.

Figures

Fig. 8. Shells of Durgella species. A: D. rhaphiellus syntypes ZMB/MOLL 5033. B–D: D. siamensis; B: syntypes SMF 227168/1; C, D: CUMZ 14232.

Fig. 1. Map of Thailand and Myanmar showing the geographic distribution of eight species of Durgella based on the specimens examined herein. Only D. rhaphiellus is not shown on the map, as its type locality of “Siam” [Thailand] is too imprecise.

Fig. 3. Living snails of seven Durgella species. A: D. birmanica CUMZ 14233. B, C: D. levicula; B: CUMZ 14248; C: CUMZ 14250. D: D. erratica CUMZ 14234. E: D. siamensis CUMZ 14232. F: D. libas CUMZ 14237. G: D. pentata sp. nov. paratype CUMZ 14240. H: D. nulla sp. nov. paratype CUMZ 14228.

Fig. 9. Microsculpture of Durgella shells: protoconch and early teleoconch (first column), and close-up view of protoconch (second column). A, B: D. birmanica CUMZ 14233. C, D: D. levicula CUMZ 14251. E, F: D. erratica CUMZ 14234. G, H: D. siamensis CUMZ 14232.

Fig. 10. Genitalia of Durgella species. A: D. erratica CUMZ 14234. B, C: D. siamensis CUMZ 14232; B: general view of the genital system; C: internal structure of the penis and epiphallus. White arrow indicates the end of the penis.

Fig. 11. Representative SEM images of the radula. A, B: D. birmanica CUMZ 14233. C, D: D. levicula CUMZ 14253. E, F: D. erratica CUMZ 14234. G, H: D. siamensis CUMZ 14232. Yellow arrow indicates the central tooth.

Fig. 6. Genitalia of Durgella species. A: D. birmanica CUMZ 14233. B, C: D. levicula CUMZ 14253; B: general view of the genital system; C internal structure of the penis and epiphallus. White arrow indicates the end of the penis.

Fig. 4. Eggs and mating behavior of Durgella species recorded in the field.A: eggs of D. libas from Wachirathan Waterfall, Chiang Mai Province. B: eggs of D. pentata sp. nov. from Tham Chiang Dao, Chiang Mai Province. C: eggs of D. nulla sp. nov. from Phu Pha Lom, Loei Province. D: mating pairs of D. nulla sp. nov. from Phu Pha Lom, Loei Province.

Fig. 12. Shells of Durgella species. A, B: D. libas CUMZ 14236. C, D: D. pentata sp. nov.; C: holotype CUMZ 14239; D: paratype CUMZ 14240. E, F: D. nulla sp. nov.; E: holotype CUMZ 14227; F: paratype CUMZ 14228.

Fig. 13. Microsculpture of Durgella shells: protoconch and early teleoconch (first column), and close-up view of protoconch (second column). A, B: D. libas CUMZ 14236. C, D: D. pentata sp. nov. paratype CUMZ 14240. E, F: D. nulla sp. nov. paratype CUMZ 14228.

Fig. 14. Genitalia of Durgella libas CUMZ 14236. A: general view of the genital system and B: internal structure of the penis and epiphallus. White arrow indicates the end of the penis.

Fig. 17. Representative SEM images of the radula. A, B: D. libas CUMZ 14238. C, D: D. pentata sp. nov. paratype CUMZ 14240. E, F: D. nulla sp. nov. paratype CUMZ 14228. Yellow arrow indicates the central tooth.

Fig. 15. Genitalia of Durgella pentata sp. nov. paratype CUMZ 14240. A: general view of the genital system and B: internal structure of the penis and epiphallus. White arrow indicates the end of the penis.

Fig. 16. Genitalia of Durgella nulla sp. nov. paratype CUMZ 14228. A: general view of the genital system and B: internal structure of the penis and epiphallus. White arrow indicates the end of the penis.