taxonID	type	description	language	source
426CF52FA4674B5FFF1B8191671FBF83.taxon	type_taxon	Type species. Nanina (Macrochlamys) diadema Dall, 1897	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4674B5FFF1B8191671FBF83.taxon	etymology	Etymology. Taphrenalla is derived from ‘ Taphr ’ (the Greek word ‘ Taphros ’) meaning ‘ trench or ditch’ and ‘ enalla ’ (the Greek word ‘ enallax ’) meaning crosswise. Therefore, the generic name means the snail with transverse ditches or radial grooves on the shell sculpture. The gender of the new generic name is feminine.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4674B5FFF1B8191671FBF83.taxon	description	Description Shell. Shell dextral, depressed to globosely depressed, thin, translucent, corneous to dark brown colour. Shell surface shiny, smooth (fig. 4 A), with very fine growth lines or sculptured, very weak (fig. 4 B) to strong (figs 4 C, D) radial grooves starting from suture. Suture shallow impressed (figs 4 A, B), deep impressed (fig. 4 C), or wide and deep channel-shaped (fig. 4 D). Whorls 5 – 6, increasing regularly. Last whorl rounded to slightly shouldered. Aperture crescentic with simple lip. Columellar margin simple and little expanded near umbilicus. Umbilicus perforate, narrowly opened. Genital organs. Genitalia with short to long penis; thick or thin penial sheath. Epiphallic caecum usually straight, rarely a little bent at tip (not spirally coiled); penial retractor muscle attached to head of epiphallic caecum (rarely attached near the head). Flagellum relatively short and enlarged at base. Penial verge well-developed and small to large; penial caecum usually present and short or rarely absent. Female genital organ well-developed; dart apparatus located on vagina or at vagina and penis junction. Spermatophore long, needle-shaped and translucent. Head filament smooth, shorter than sperm sac and gradually thinning to terminal point. Sperm sac enlarged, capsule-shaped and contained whitish sperm mass. Tail filament very long and thin tube; region close to sperm sac bearing one or two spines. Radula. Teeth arranged in U-shape with very wide angle. Central tooth symmetrical tricuspid; mesocone lanceolated-shaped; ectocones short and located at middle of tooth height. Lateral teeth asymmetrical tricuspid; mesocone triangular shaped; endocone smaller than ectocone; ectocone located below endocone. Marginal teeth with obliquely bicuspid, elongate and narrower teeth, and outermost teeth shorter and smaller than inner teeth (fig. 4 H). External features. Living snails usually covered with reticulated skin; three colourful stripes running from head to tail (one at middle of body and two on each side of body near foot sole). Eye stalks and head usually with dark colour; tail region usually with lighter colour than head region. Pallial cavity and some internal organs visible through translucent shell (figs 4 F, G). Mantle collar large, well developed and composed of two shell lobes and three dorsal lobes (figs 4 E – G); shell lobes usually slender and longer than dorsal lobes. Right shell lobe placed on upper shell surface and longer than left shell lobe placed below periphery. Right dorsal lobe larger than left dorsal lobe; left dorsal lobe divided into anterior left dorsal lobe and posterior left dorsal lobe. Foot sole tripartite; caudal foss long and narrow; caudal horn raised and large. Colouration of soft body usually variable from reddish, orange, yellow, milky-white to greyish, and colour disappeared in preserved specimens.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4674B5FFF1B8191671FBF83.taxon	distribution	Distribution range. Endemic to Malay Peninsula: Southern Thailand (fig. 1) and probably in Peninsular Malaysia.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4674B5FFF1B8191671FBF83.taxon	discussion	Remarks. Taphrenalla diadema is the type species of the new genus because this species somewhat represents the morphology in general among congeneric species and is also the oldest valid species. Although the absence of radial grooves is evolved in T. zemia sp. nov. and T. alba sp. nov., they present the other characteristics of yellowish stripes on the body and a simple spermatophore. The spermatophore of T. alba sp. nov. has two spines near the sperm sac but is spineless in other regions. In addition, the molecular analysis grouped them within the Taphrenalla clade with strong support. For comparison with the other closely related ariophantid genera, Taphrenalla gen. nov. can be distinguished from Sarika, Macrochlamys, Taphrospira, Syama, and Rasama by the shell sculpture having radial grooves, while the others have a smooth shell without radial grooves (table 5). The shell morphology of Hemiplecta and Cryptozona are clearly different from the new genus in being much larger, with a smooth shell surface and without a radial groove. Regarding the genitalia anatomy, Taphrenalla gen. nov. have a penial verge, dart apparatus, and straight or slightly bent epiphallic caecum. Compared to the other genera, Macrochlamys and Syama both have a spirally coiled epiphallic caecum, but only Syama has no dart apparatus. The genus Taphrospira also has a straight epiphallic caecum and is without a dart apparatus, while the genus Rasama has neither an epiphallic caecum nor a dart apparatus (Blanford & Godwin-Austen, 1908; Schileyko, 2002, 2003). Thus, the genitalia of the genus Sarika is similar to Taphrenalla gen. nov. However, all species of the genus Sarika, except for S. consepta (Benson, 1860), have no penial verge (Pholyotha et al., 2020), while this character is present in all Taphrenalla species. The taxonomic status of S. consepta is needed to be investigated in the future using a molecular approach. In addition, most Taphrenalla species have bright body colouration patterns of the left, middle, and right stripes, either restricted to the head section or running from head to tail, while all Sarika species have a monochrome greyish to blackish body colour (Pholyotha et al., 2020).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4614B5DFD3E81D76132BAB8.taxon	description	(figs 1, 3, 5, 6 A, 7 A – C, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4614B5DFD3E81D76132BAB8.taxon	materials_examined	Material examined. Type material: not examined. Non-type material: Wat Tham Wararam, Phanom District, Surat Thani (98 ° 53 ’ 07.3 ” N 98 ° 40 ’ 01.0 ” E): CUMZ 7153 (fig. 6 A; 26 shells and 15 specimens in ethanol), CUMZ 7154 (six specimens in ethanol), CUMZ 7155 (14 shells and eight specimens in ethanol), and CUMZ 7156 (four shells). Limestone outcrops at Khlong Sok, Phanom District, Surat Thani (8 ° 53 ’ 39.3 ” N 98 ° 33 ’ 10.7 ” E): CUMZ 7157 (three shells) and CUMZ 7158 (one shell). Limestone outcrops at Khlong Sok, Phanom District, Surat Thani (8 ° 50 ’ 51.2 ” N 98 ° 44 ’ 18.0 ” E): CUMZ 7159 (31 shells and 12 specimens in ethanol). Wat Tham Phanthurat, Phanom District, Surat Thani (8 ° 54 ’ 01.3 ” N 98 ° 31 ’ 30.2 ” E): CUMZ 7160 (three specimens in ethanol).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4614B5DFD3E81D76132BAB8.taxon	discussion	Remarks. Taphrenalla asamurai had been re-described and illustrated by Sutcharit & Panha (2008). The unique characters are a depressed conic shell of medium size (width up to 21.1 mm; height up to 11.7 mm), deep and narrow impressed suture, and long radial grooves that appeared on the last 37 – 64 grooves (n = 32; figs 5, 6 A). The genitalia are bent at the epiphallic caecum, and the proximal part of dart apparatus is with a thick and swollen sac-like structure (figs 7 A, 7 B). Spermatophore has one spine on the tail filament close to the sperm sac (fig. 7 C). Living snails have monochrome red to dark red on tail (fig. 3).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	description	(figs 1, 3, 5, 6 B, 7 D – F, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	materials_examined	Type material examined. Holotype: CUMZ 7161 (fig. 6 B). Paratypes. Same data as holotype: CUMZ 7162 (15 shells and five specimens in ethanol), CUMZ 7163 (three shells and nine specimens in ethanol), NHMUK (two shells) and SMF (two shells). Type locality. Limestone outcrops at Khlong Sok, Phanom District, Surat Thani, Thailand (8 ° 53 ’ 20.5 ” N 98 ° 40 ’ 27.7 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	etymology	Etymology. The specific epithet is derived from ‘ macro ’ (the Greek word ‘ makros ’) meaning ‘ long’ and ‘ sulcata ’ (the Latin word ‘ sulcus ’) meaning ‘ furrow, groove’ with reference to the characteristic of having numerous long radial grooves on shell sculpture.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	diagnosis	Diagnosis. Shell with a narrow and deep impressed suture and deep and long radial grooves. Epiphallic caecum straight and short. Penial caecum short. Inner sculpture of proximal penis with fine wrinkled longitudinal folds; distally with oblique longitudinal folds and transverse-folded surrounding penial verge. Dart apparatus located on vagina. Spermatophore with one spine on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	description	Description Shell (fig. 6 B). Shell comparatively depressed conic, medium-sized (width up to 18.7 mm; height up to 9.8 mm) and pale brown. Whorls 5 – 6; suture deep and narrow impressed. Upper shell surface with strong and long radial grooves continuing from suture, gradually inconspicuous and then disappear below periphery. Radial grooves starting between whorl 3 to 4 and with 42 – 58 grooves at last whorl (n = 24; fig. 5). Spire depressed conic; apex raised with a spire angle of about 143 – 164 °. Last whorl shouldered and well-rounded at periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (figs 7 D – F). Atrium short. Penis elongate cylindrical and same length as total vagina length; penial caecum present. Inner sculpture of proximal penial wall has very fine wrinkled, longitudinal penial pilasters, with thick and oblique longitudinal folds in middle, and then transformed to a thin, transverse-folded, surrounded penial verge. Epiphallus slender tube and about two times total penis length. Epiphallic caecum very short; thick penial retractor muscle attached at tip. Flagellum large and irregular slender tube. Vas deferens thin tube. Vagina long and cylindrical tube. Dart apparatus located on vagina away from penis joining to atrium. Spermatophore long and needle-shaped. Tail filament very long tube; region close to sperm sac bearing a very small spine. External features (fig. 3). Animal dark gray mixed with pale yellow on head and body and monochrome bright yellowish on tail.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4634B63FF1B85126725BC37.taxon	discussion	Remarks. Taphrenalla macrosulcata sp. nov. is similar to T. asamurai in shell morphology but can be differentiated by a paler brown shell and a pale yellowish soft body. Whereas, T. asamurai has a dark brown shell and reddish to dark red soft body. Regarding the genital anatomy, this new species has a short flagellum, straight epiphallic caecum, and dart apparatus connecting to the vagina away from the penis junction and without thick connective tissue surrounding the proximal end. For comparison, T. asamurai has a long flagellum, bent epiphallic caecum, and dart apparatus located opposite the penis junction and with thick connective tissue at the proximal end. The presence of one spine on the tail filament of the spermatophore is the remarkable character of T. macrosulcata sp. nov. and T. asamurai, while other Taphrenalla species have two spines.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45D4B60FD3E829B676DBF4F.taxon	description	(figs 1, 3, 5, 8 A – C, 9 A – C, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45D4B60FD3E829B676DBF4F.taxon	materials_examined	Material examined. Type material: holotype USNM 150277. Non-type material: Wat Khuha Santayaram (Wat Tham Khao Daeng), Ron Phibun District, Nakhon Si Thammarat (8 ° 14 ’ 39.5 ” N 99 ° 52 ’ 00.1 ” E): CUMZ 7164 (one specimen in ethanol). Tham Talot, Thung Song District, Nakhon Si Thammarat (8 ° 09 ’ 32.3 ” N 99 ° 40 ’ 41.6 ” E): CUMZ 7165 (16 shells and three specimens in ethanol). Tham Rad, Thung Song District, Nakhon Si Thammarat (8 ° 02 ’ 47.2 ” N 99 ° 43 ’ 45.0 ” E): CUMZ 7166 (10 shells) and CUMZ 7167 (11 specimens in ethanol). Wat Tham Phraphut, Ratsada District, Trang (7 ° 57 ’ 57.3 ” N 99 ° 44 ’ 41.3 ” E): CUMZ 7168 (four shells and two specimens in ethanol). Tham Chang Hai, Mueang District, Trang (7 ° 35 ’ 26.6 ” N 99 ° 40 ’ 05.3 ” E): CUMZ 7169 (nine shells) and CUMZ 7170 (two shells). Khao Pu Chao Bureau of Monks, Na Yong District, Trang (7 ° 33 ’ 53.0 ” N 99 ° 46 ’ 36.2 ” E): CUMZ 7171 (fig. 8 C; 18 shells) and CUMZ 7172 (four shells). Wat Khao Huai Haeng, Huai Yot District, Trang (7 ° 47 ’ 36.5 ” N 99 ° 38 ’ 37.0 ” E): CUMZ 7173 (16 specimens in ethanol). Wat Tham Sumano, Srinagarindra District, Phatthalung (7 ° 35 ’ 14.0 ” N 99 ° 52 ’ 06.8 ” E): CUMZ 7174 (10 specimens in ethanol) and CUMZ 7175 (fig. 8 A; four shells and 33 specimens in ethanol). Khao Ok Talu Khuha Sawan, Mueang District, Phatthalung (7 ° 37 ’ 33.7 ” N 100 ° 05 ’ 23.7 ” E): CUMZ 7176 (three shells) and CUMZ 7177 (fig. 8 B; 55 shells and 18 specimens in ethanol). Wat Khuha Sawan, Mueang District, Phatthalung (7 ° 37 ’ 15.0 ” N 100 ° 04 ’ 48.7 ” E): CUMZ 7178 (one shell). Tham Phaya Hong, Kong Ra District, Phatthalung (7 ° 27 ’ 48.4 ” N 99 ° 57 ’ 53.3 ” E): CUMZ 7179 (36 shells) and CUMZ 7180 (22 shells and 137 specimens in ethanol). Limestone outcrops at Khlong Chaloem, Kong Ra District, Phatthalung (7 ° 23 ’ 50.6 ” N 99 ° 58 ’ 45.2 ” E): CUMZ 7181 (five shells and 31 specimens in ethanol). Tham Phra Non, Mueang District, Phatthalung (7 ° 40 ’ 52.1 ” N 100 ° 03 ’ 42.4 ” E): CUMZ 7182 (10 shells).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45D4B60FD3E829B676DBF4F.taxon	discussion	Remarks. Taphrenalla diadema had been recently re-described and illustrated by Sutcharit & Panha (2008). The distinguishing characters are a depressed conic shell, medium-sized (width up to 17.8 mm; height up to 10.6 mm), a wide and deep channel-shaped suture, and short radial grooves appeared on last about 24 – 43 grooves (n = 39; figs 5, 8 A – C). The genitalia have a short and straight epiphallic caecum, and dart apparatus located on vagina away from vagina and penis junction (figs 9 A, B). Spermatophore has one short spine on tail filament close to sperm sac (fig. 9 C). Living snail has yellowish stripes on head and tail (fig. 3). The specimens from Khao Ok Talu, Phatthalung Province (fig. 8 B) tended to have a smaller diameter and shallower radial grooves than the other populations (figs 8 A, C). However, all the genital characters and spermatophores from all localities appear to be identical. In addition, the molecular data also confirmed that the shell variations were all grouped within the T. diadema clade.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	description	(figs 1, 3, 5, 8 D, E, 9 D – F, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	materials_examined	Type material examined. Holotype: CUMZ 7183 (fig. 8 D). Paratypes. Same data as holotype: CUMZ 7184 (31 shells and nine specimens in ethanol), CUMZ 7188 (one shell), NHMUK (two shells) and SMF (two shells). Other material examined. Wat Khiri Rat Phatthana, Wiang Sa District, Surat Thani (8 ° 31 ’ 36.5 ” N 99 ° 23 ’ 00.2 ” E): CUMZ 7185 (63 shells and 88 specimens in ethanol). Wat Na San, Ban Na San District, Surat Thani, Thailand (8 ° 48 ’ 30.7 ” N 99 ° 22 ’ 16.1 ” E): CUMZ 7186 (four shells and 11 specimens in ethanol) and CUMZ 7187 (11 shells and 24 specimens in ethanol). Wat Tham Kanlayanamit, Tham Phannara District, Nakhon Si Thammarat (8 ° 30 ’ 49.3 ” N 99 ° 22 ’ 53.9 ” E): CUMZ 7189 (fig 8 E; 39 shells and 27 specimens in ethanol). Wat Tham Thong Panara, Tham Phannara District, Nakhon Si Thammarat (8 ° 25 ’ 19.1 ” N 99 ° 22 ’ 48.4 ” E): CUMZ 7190 (20 shells and one specimen in ethanol). Type locality. Limestone outcrop at Tham Lord, Nopphitam District, Nakhon Si Thammarat, Thailand (8 ° 47 ’ 16.7 ” N 99 ° 38 ’ 29.9 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	etymology	Etymology. This species is named after William Healey Dall, an American malacologist, who discovered the first species of Crown Snail, T. diadema.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	diagnosis	Diagnosis. Shell with a very wide channel-shaped suture and with deep and short radial grooves. Epiphallic caecum straight. Penial caecum very short. Inner sculpture of proximal penis with very small, irregular and longitudinal folds, and then with obliquely and wrinkled to very fine pilasters distally. Dart apparatus connected on vagina. Spermatophore with two spines on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	description	Description Shell (figs 8 D, E). Shell depressed conic to globosely depressed, medium-sized (width up to 18.3 mm; height up to 11.3 mm) and pale to dark brown. Whorls 5 – 6; suture very wide, deep and channeled-shape. Upper shell surface with strong and short radial grooves continuing from suture to inconspicuous at periphery and then disappear below periphery. Radial grooves beginning between whorl 3 to 4 and having 23 – 37 grooves at last whorl (n = 33; fig. 5). Spire depressed conic; apex raised with a spire angle of about 135 – 155 °. Last whorl shouldered and well-rounded at periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (figs 9 D – F). Atrium slightly short. Penis elongate cylindrical and nearly two times that of total vagina length; penial caecum present. Inner sculpture of proximal penial wall supported by penial pilasters starting proximally with very small irregular longitudinal folds, then gradually, obliquely, wrinkled pilasters in middle, to very fine pilasters surrounding penial verge. Epiphallus slender tube, approximately one and a half times total penis length. Epiphallic caecum slightly prolonged, and attached with thick penial retractor muscle (prm). Flagellum enlarged at base and tapering with irregular shape to tip. Vas deferens slender tube. Vagina long and cylindrical tube. Dart apparatus located on proximal vagina away from penis joining to atrium. Spermatophore long and needle-shaped. Tail filament very long tube; region close to sperm sac bearing two spines; spine close to sperm sac shorter than the other. External features (fig. 3). Animal dark gray in body colour with three pale orange stripes running from head to caudal horn. One stripe at middle of the body and others two stripes on side of body close to foot sole; stripes on tail wider than anterior.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45E4B67FD1F8104618ABD08.taxon	discussion	Remarks. Shell morphology of this new species looks similar to T. diadema, but can be distinguished by having a wider base of the suture than in T. diadema. In comparison, T. dalli sp. nov. shows a small penial caecum, shorter flagellum, and large and coarse penial sculpture pilaster at the middle of penis. Whereas, T. diadema has a large penial caecum, longer flagellum, and inner wall of penis is sculptured with very fine wrinkled pilasters. In addition, the penial sculpture at the middle of the penis of T. dalli sp. nov. has larger, more obliquely wrinkled pilasters than T. diadema. This new species shows variation in size and shell sculpture, the like of which also appears in T. diadema. However, the shell suture and genitalia of all variants are identical, and are also clustered together in the T. dalli clade in the molecular analysis.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	description	(figs 1, 3, 5, 10 A, 11 A – C, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	materials_examined	Type material examined. Holotype: CUMZ 7191 (fig. 10 A). Paratypes. Same data as holotype: CUMZ 7192 (two shells and nine specimens in ethanol), CUMZ 7193 (21 shells), CUMZ 7194 (one specimen in ethanol), NHMUK (two shells), and SMF (two shells). Other material examined. Limestone outcrops at Ao Luek Tai, Ao Luek District, Krabi (8 ° 24 ’ 09.9 ” N, 98 ° 44 ’ 16.3 ” E): CUMZ 7195 (five specimens in ethanol). Tham Sa Yuan Thong, Ao Luek District, Krabi (8 ° 21 ’ 44.6 ” N, 98 ° 44 ’ 45.9 ” E): CUMZ 7196 (one shell). Tham Phet, Ao Luek District, Krabi (8 ° 23 ’ 33.9 ” N, 98 ° 46 ’ 24.0 ” E): CUMZ 7197 (one shell). Wat Khao Hua Sing, Plai Phraya District, Krabi (8 ° 30 ’ 47.7 ” N 98 ° 45 ’ 30.1 ” E): CUMZ 7198 (29 shells). Tham Nalakiring Bureau of Monks, Plai Phraya District, Krabi (8 ° 33 ’ 35.3 ” N 98 ° 51 ’ 37.5 ” E): CUMZ 7225 (eight shells and three specimens in ethanol). Wat Khao Phra, Phrasaeng District, Surat Thani (8 ° 37 ’ 33.0 ” N 98 ° 56 ’ 53.5 ” E): CUMZ 7226 (15 shells). Tham Namlod Bureau of Monks, Phrasaeng District, Surat Thani (8 ° 40 ’ 40.2 ” N 98 ° 56 ’ 41.9 ” E): CUMZ 7227 (six shells). Figure 10 Shells. A. Taphrenalla corona sp. nov. holotype CUMZ 7191. B. T. parversa sp. nov. holotype CUMZ 7199. C. T. conformis sp. nov. holotype CUMZ 7202. D. Taphrenalla incilis sp. nov. holotype CUMZ 7207. Inset SEM image showing shell sculpture of each species. Downloaded from Brill. com 10 / 07 / 2022 07: 10: 01 PM via free access Figure 11 Genitalia and spermatophore. A – C. Taphrenalla corona sp. nov. paratype CUMZ 7192. A. whole reproductive system. B. internal wall of penis with penial verge. C. spermatophore; inset showing two spines located close to the sperm sac. D – F. T. parversa sp. nov. paratype CUMZ 7200. D. whole reproductive system. E. internal wall of penis with penial verge. F. spermatophore; inset showing two spines located close to the sperm sac. White arrowheads indicate the position of penial verge. Type locality. Limestone outcrops near Than Bok Khorani, Ao Luek District, Krabi, Thailand (8 ° 22 ’ 28.7 ” N 98 ° 44 ’ 07.9 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	etymology	Etymology. The specific epithet “ corona ” is from the Latin word “ corona ” meaning “ crown ” and is in reference to the characteristic radial grooves on the shell sculpture of this genus.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	diagnosis	Diagnosis. Shell with wide channel-shaped suture, and slightly deep and short radial grooves. Penis with thick penial sheath. Penial caecum absent. Epiphallic caecum straight. Inner sculpture of proximal penis with fine irregular longitudinal folds and then with obliquely longitudinal folds distally. Dart apparatus connected on vagina. Spermatophore with two spines on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	description	Description Shell (fig. 10 A). Shell depressed conic, medium-sized (width up to 16.0 mm; height up to 9.3 mm) and slightly milky to brownish. Whorl 5 – 6; suture wide and deep groove, channel-shaped. Upper shell surface with rather strong and short radial grooves continuing from suture to inconspicuous at periphery and disappears below periphery. Radial grooves starting between whorl 3 to 4 and having 26 – 47 grooves at last whorl (n = 26; fig. 5). Spire slightly depressed conic; apex raised with a spire angle of about 140 – 155 °. Last whorl barely shouldered and well-rounded at periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (figs 11 A – C). Atrium short. Penis enlarge cylindrical and same length as vagina; penial caecum absent. Inner sculpture of proximal penis with fine irregularly longitudinal penial pilasters, gradually transforming to obliquely longitudinal folds surrounding penial verge. Epiphallus slender tube, about two and a half times total penis length and smaller diameter than penis. Epiphallic caecum short, and attached with thick penial retractor muscle. Flagellum prolonged with irregular cylindrical tip. Vas deferens slim tube. Vagina prolonged, cylindrical-shaped and distal part larger diameter than proximal part. Dart apparatus located on proximal vagina away from penis joining to atrium. Spermatophore long and needle-shaped. Tail filament very long tube; region near sperm sac bearing two spines, spine close to sperm sac very small and shorter than the other. External features (fig. 3). Animal very dark gray in body colour with one creamy to pale orange stripe in middle of body running from head to caudal horn; stripe on tail wider than on body to head.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4594B64FD3E83426022BA29.taxon	discussion	Remarks. Although the shell morphology of this new species is close to those of T. diadema and T. dalli sp. nov., its genitalia is unique. The absence of a penial caecum and presence of a thick penial sheath covering a proximal portion of penis distinguished T. corona sp. nov. from the other Taphrenalla species.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	description	(figs 1, 3, 10 B, 11 D – F, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	materials_examined	Type material examined. Holotype: CUMZ 7199 (fig. 10 B). Paratypes. Same data as holotype: CUMZ 7200 (one shell and five specimens in ethanol), CUMZ 7201 (five shells), NHMUK (two shells), and SMF (two shells). Type locality. Tham Suwan Khuha, Takua Thung District, Phang-nga, Thailand (8 ° 25 ’ 45.0 ” N, 98 ° 28 ’ 19.6 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	etymology	Etymology. This species is derived from ‘ parv ’ (the Latin word ‘ parvus ’) meaning ‘ little’ and ‘ versa ’ (the Latin word ‘ versus ’) meaning ‘ line, furrow’ with reference to the characteristic numerous little radial grooves on the shell sculpture.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	diagnosis	Diagnosis. Shell with shallowly impressed suture and very weakly radial grooves. Penis with thick penial sheath. Penial caecum absent. Epiphallic caecum straight. Inner sculpture of proximal penis with fine, irregular and longitudinal folds, and then trasition to irregular and zig-zag, longitudinal folds distally. Dart apparatus connected on vagina. Spermatophore with two spines on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	description	Description Shell (fig. 10 B). Shell comparatively depressed conic, medium-sized (width up to 14.4 mm; height up to 8.1 mm) and pale brown. Whorls 5 – 6; suture shallowly impressed. Upper shell surface with numerous weak radial grooves continuing from suture and disappear before periphery. Spire depressed conic; apex raised with a spire angle of about 142 – 164 °. Last whorl barely shouldered and rounded periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (figs 11 D – F). Atrium very short. Penis stout cylindrical and about same length as vagina; penial caecum absent. Inner sculpture of proximal penial wall with fine irregular longitudinal penial pilasters and then gradually irregular, zig-zag, longitudinal folds in middle and surrounded penial verge. Epiphallus slender tube, around two times total penis length, and smaller diameter than penis; proximal epiphallus larger than distal portion. Epiphallic caecum short, same diameter as proximal epiphallus, and attached with thick penial retractor muscle. Flagellum short with irregular shape. Vas deferens small tube. Vagina very enlarged, cylindrical-shaped, and distal part enlarged. Dart apparatus located on proximal vagina away from penis joining to atrium. Spermatophore elongate and needle-shaped. Tail filament very long tube; region close to sperm sac bearing two spines; spine located close to sperm sac smaller and shorter than the other. External features (fig. 3). Animal dark gray colour on head to body with three bright orange stripes running from head to body (two stripes near foot sole and one at middle of body), and then gradually only bright orange stripes on body to tail.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA45A4B6BFD1F84A1604EBD99.taxon	discussion	Remarks. Taphrenalla parversa sp. nov. is similar to T. corona sp. nov. because of the absence of the penial caecum and a thick penial sheath. However, they can be discriminated by their flagellum, epiphallus, vagina, and shell morphology. Taphrenalla parversa sp. nov. has a shorter flagellum, longer epiphallus, and larger vagina than T. corona sp. nov. In shell comparison, T. parversa sp. nov. has a weakly impressed suture and very weakly radial grooves on the shell sculpture, whereas T. corona sp. nov. exhibits a wide and deep groove at suture and deep radial grooves on the shell sculpture.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	description	(figs 1, 3, 10 C, 12 A – C, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	materials_examined	Type material examined. Holotype: CUMZ 7202 (fig. 10 C). Paratypes. Same data as holotype: CUMZ 7203 (15 shells and 52 specimens in ethanol), CUMZ 7230 (three specimens in ethanol), CUMZ 7204 (eight specimens in ethanol), CUMZ 7205 (11 shells and six specimens in ethanol), CUMZ 7206 (24 specimens in ethanol), NHMUK (two shells), and SMF (two shells). Type locality. Tham Phung Chang, Mueang District, Phang-nga, Thailand (8 ° 26 ’ 33.0 ” N, 98 ° 30 ’ 55.4 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	etymology	Etymology. The specific name is from the Latin ‘ conformis ’ meaning “ like ” or “ similar ” and name refers to the new species possessing a superficially similar shell to T. parversa sp. nov.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	diagnosis	Diagnosis. Shell having shallowly impressed suture, weakly radial grooves. Penis covered with thick penial sheath. Epiphallic caecum short and bent at tip. Penial caecum short. Inner sculpture of proximal penis with very small longitudinal folds; distally narrow and obliquely cuboidal folds. Dart apparatus connected on vagina. Spermatophore with two spines on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	description	Description Shell (fig. 10 C). Shell comparatively depressed conic, medium-sized (width up to 17.7 mm; height up to 10.3 mm) and pale brown. Whorls 5 – 6; suture shallowly impressed. Upper shell surface with numerous weak radial grooves continuing from suture to gradually disappear before periphery. Spire depressed conic; apex raised with a spire angle of about 138 – 148 °. Last whorl well-rounded periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (figs 12 A – C). Atrium slightly prolonged. Penis cylindrical and nearly three times total vagina length; penial caecum present. Inner sculpture of proximal penis with small longitudinal folds; distal portion with narrow obliquely cuboidal pilasters, then reduced in size and surrounding penial verge. Epiphallus slender tube, around two times total penis length and smaller diameter than penis. Epiphallic caecum short and bent at the tip; thick penial retractor muscle connecting near the tip. Flagellum short, tapering and irregular slender tube. Vas deferens thin tube. Vagina long and cylindrical-shaped. Dart apparatus located near proximal vagina away from penis join to atrium. Spermatophore long and needle-shaped. Tail filament very long tube; region close to sperm sac bearing two spines; spine close to sperm sac small and shorter than the other. External features (fig. 3). Animal dark gray in body colour with three pale milky yellow stripes running from head to caudal horn. One stripe at middle of the body and other two stripes above foot sole.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4554B69FD3E843267FFBC30.taxon	discussion	Remarks. Shell characters in T. conformis sp. nov. and T. parversa sp. nov. are very similar. They have a weakly impressed suture and weakly radial grooves on shell sculpture. However, the body colouration and reproductive organ can be used to separate them. In T. conformis sp. nov., the body has three pale milky-yellow bands running from the head to caudal horn, and the genitalia have a penial caecum. Whereas, the body colouration of T. parversa sp. nov. contains three bright orange stripes running from head to body, and only bright orange stripes on the body to tail. The genitalia of T. parversa sp. nov. do not have a penial caecum.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4574B68FD3E829B67F3BF83.taxon	description	(figs 1, 3, 10 D, 12 D – F, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4574B68FD3E829B67F3BF83.taxon	materials_examined	Type material examined. Holotype: CUMZ 7207 (fig. 10 D). Paratypes. Same data as holotype: CUMZ 7208 (four shells and five specimens in ethanol), CUMZ 7209 (six shells and six specimens in ethanol), NHMUK (two shells), and SMF (two shells). Type locality. Tham Khao Ting, Palian District, Trang, Thailand (7 ° 09 ’ 30.0 ” N, 99 ° 48 ’ 06.0 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4574B68FD3E829B67F3BF83.taxon	etymology	Etymology. The specific epithet “ incilis ” is from the Latin word “ incile ” meaning “ ditch, trench ”, which referrs to the characteristic small radial grooves in the shell sculpture. Diagnosis. Shell with narrow and deep channel-shaped suture, and weak and short radial grooves. Epiphallic caecum short and bent at tip. Penial caecum short. Inner sculpture of proximal penis with very small longitudinal folds; distally narrow and obliquely cuboidal folds. Dart apparatus located at vagina and penis junction. Spermatophore with two spines on tail filament close to sperm sac.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4574B68FD3E829B67F3BF83.taxon	description	Description Shell (fig. 10 D). Shell comparatively depressed conic, moderate-sized (width up to 16.1 mm; height up to 8.8 mm) and pale to dark brown. Whorls 5 – 6; suture narrow, deep grooved, and channel-shaped. Upper shell surface with numerous, very short and weak radial grooves near suture and then suddenly disappear. Spire depressed conic; apex raised with a spire angle of about 134 – 151 °. Last whorl barely shouldered, and periphery almost rounded. Aperture lip simple; ummbilicus narrowly opened. Genital organs (figs 12 D – F). Atrium short. Penis cylindrical and around three times total vagina length; penial caecum present. Inner sculpture of proximal and middle part of penis with fine longitudinal folds; distal portion with narrow obliquely cuboidal penial pilasters surrounding penial verge. Epiphallus slender tube and slightly longer than penis. Epiphallic caecum short and bent at tip; thick penial retractor muscle connecting near the tip. Flagellum short, irregular curved and with obtuse tip. Vas deferens thin tube. Vagina prolonged, cylindrical-shaped. Dart apparatus located at junction of vagina and penis. Spermatophore long and needle-shaped. Tail filament very long tube; region close to sperm sac bearing two spines; spine near sperm sac very small and shorter than the other. External features (fig. 3). Animal dark gray in body colour with a pale milky yellow stripe at middle of the body running from head to caudal horn. Posterior part (body to tail) with pale milky yellow colour and wider stripe than anterior part (head to body).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4574B68FD3E829B67F3BF83.taxon	discussion	Remarks. The weak radial grooves on the upper shell surface of T. incilis sp. nov. is similar to T. parversa sp. nov., T. conformis sp. nov., and T. pygmaea sp. nov. However, together with the suture shape and genitalia, T. incilis sp. nov. have a deeper and wider suture than T. parversa sp. nov. and T. conformis sp. nov. The dart apparatus of T. parversa sp. nov. and T. conformis sp. nov. is located on the proximal vagina away from the penis junction, while T. incilis sp. nov. has the dart apparatus joining at the junction of vagina and penis. While the suture shape and position of dart apparatus of T. incilis sp. nov. is similar to T. pygmaea sp. nov., T. incilis sp. nov. have a bent epiphallic caecum, and short flagellum and vagina, whereas T. pygmaea sp. nov. have a straight epiphallic caecum, and long flagellum and vagina.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	description	(figs 1, 3, 13 A, 14 A, B, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	materials_examined	Type material examined. Holotype: CUMZ 7210 (fig. 13 A). Paratypes. Same data as holotype: CUMZ 7211 (19 shells and 12 specimens in ethanol), NHMUK (two shells), and SMF (two shells). Other material examined. Wat Sathit Khirirom, Khiri Rat Nikhom District, Surat Thani (9 ° 01 ’ 48.2 ” N 98 ° 59 ’ 12.5 ” E): CUMZ 7212 (75 shells and nine specimens in ethanol). Type locality. Tham Wang Badan Priest’s Camp Site, Khiri Rat Nikhom District, Surat Thani, Thailand (8 ° 54 ’ 21.9 ” N 98 ° 56 ’ 53.9 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	etymology	Etymology. The specific epithet ‘ pygmaea ’ is from the Latin word ‘ pygmaeus ’ meaning ‘ dwarf’, which referrs to it being the smallest sized snail in this genus.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	diagnosis	Diagnosis. Shell small to medium-sized, narrow and deep channel-shaped suture, and weak, diminutive radial grooves. Epiphallic caecum straight. Penial caecum short. Inner sculpture of proximal penis with very fine longitudinal folds and then transition to oblique and transverse folds distally. Dart apparatus located at junction of vagina and penis.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	description	Description Shell (fig. 13 A). Shell moderately depressed conic, small to nearly medium-sized (width up to 12.3 mm; height up to 6. 8 mm) and pale brown. Whorls 5 – 6; suture narrow, deep, and channel-shaped. Upper shell surface with weak and diminutive radial grooves that appear only near suture. Spire deressed conic; apex raised with a spire angle of about 131 – 151 °. Last whorl barely shouldered and periphery almost rounded. Aperture lip simple; umbilicus narrowly opened. Figure 13 Shells. A. Taphrenalla pygmaea sp. nov. holotype CUMZ 7210. B. T. zemia sp. nov. holotype CUMZ 7213. C. T. alba sp. nov. holotype CUMZ 7217. Inset SEM image showing shell sculpture of each species. Genital organs (figs 14 A, B). Atrium short. Penis funnel-shaped and slightly longer than vagina; penial caecum present. Inner sculpture of proximal penis with very fine longitudinal penial pilasters, gradually transforming to oblique and transverse folds surrounding penial verge. Epiphallus cylindrical-shaped, about one and half times total penis length. Epiphallic caecum short, and attached with thin penial retractor muscle. Flagellum prolonged and cylindrical with blunt tip. Vas deferens small tube. Vagina long and cylindrical-shaped. Dart apparatus located at junction of proximal vagina and proximal penis. Figure 14 Genitalia. A, B. Taphrenalla pygmaea sp. nov. paratype CUMZ 7211. A. whole reproductive system. B. internal wall of penis with penial verge. C. T. zemia sp. nov. paratype CUMZ 7214, whole reproductive system from juvenile specimen. External features (fig. 3). Animal very dark gray in body colour with indistinct pale milky stripe at middle of the body running from head to caudal horn.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4564B6DFD1F81D763ADBF72.taxon	discussion	Remarks. Taphrenalla pygmaea sp. nov. is distinctive in being the smallest of the known Taphrenalla species. Compared to T. incilis sp. nov., this new species is distinguished by a small shell with narrow body whorl. Moreover, the internal distal penial surface of T. pygmaea sp. nov. has oblique and transverse folds whereas the distal portion of T. incilis sp. nov. has narrow obliquely cuboidal pilasters.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	description	(figs 1, 3, 13 B, 14 C, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	materials_examined	Type material examined. Holotype: CUMZ 7213 (fig. 13 B). Paratypes. Same data as holotype: CUMZ 7214 (25 shells and 30 specimens in ethanol), NHMUK (two shells), and SMF (two shells). Other material examined. Tham Pha Sua Bureau of Monks, Mueang District, Phang-nga (8 ° 28 ’ 30.9 ” N 98 ° 32 ’ 20.8 ” E): CUMZ 7215 (five specimens in ethanol). Wat Tham Bang Toei, Mueang District, Phang-nga (8 ° 27 ’ 47.9 ” N 98 ° 34 ’ 11.6 ” E): CUMZ 7216 (five specimens in ethanol). Type locality. Wat Khiriwong, Thap Put District, Phang-nga, Thailand (8 ° 31 ’ 53.2 ” N 98 ° 34 ’ 41.4 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	etymology	Etymology. The specific epithet is from the Greek word ‘ zemia ’ meaning ‘ loss’, which referrs to the shell being without radial grooves on the upper surface.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	diagnosis	Diagnosis. Shell moderate to large-sized, shallowly impressed suture, and without radial groove. Epiphallic caecum straight. Dart apparatus located on vagina.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	description	Description Shell (fig. 13 B). Shell globosely depressed, moderate to nearly large (width up to 21.3 mm; height up to 11.9 mm) and yellowish-brown. Whorls 5 – 6; suture shallowly impressed. Upper shell surface smooth without radial grooves. Spire moderately elevated; apex raised with a spire angle of about 137 – 151 °. Last whorl slightly shouldered and almost rounded periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (fig. 14 C). Atrium long. Penis slender tube and slightly longer than vagina. Epiphallus cylindrical tube, nearly one and a half times total penis length. Epiphallic caecum short, and attached with penial retractor muscle. Flagellum large and slender tube. Vas deferens thin tube connected between distal end of epiphallus and free oviduct. Vagina long and cylindrical-shaped. Dart apparatus located near proximal vagina away from penis joining to atrium. External features (fig. 3). Animal dark gray in body colour with three pale milky yellow stripes running from head to caudal horn. One stripe at middle of the body and two stripes close to foot sole.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B6DFF1B81586759BD98.taxon	discussion	Remarks. Taphrenalla zemia sp. nov. is distinguished from all other Taphrenalla species by the shell having a shallowly impressed suture without a subsutural band and no radial grooves on shell sculpture. Although we surveyed three localities during the wet season, only juvenile specimens were found for the examination and illustrated herein.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	description	(figs 1, 3, 13 C, 15, table 6)	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	materials_examined	Type material examined. Holotype: CUMZ 7217 (fig. 13 C). Paratypes. Same data as holotype: CUMZ 7218 (12 shells and 22 specimens in ethanol), CUMZ 7219 (13 specimens in ethanol), NHMUK (two shells), and SMF (two shells). Other material examined. Limestone outcrops at Khlong Thom Nuea, Khlong Thom District, Krabi (7 ° 55 ’ 47.9 ” N 99 ° 16 ’ 23.3 ” E): CUMZ 7220 (six shells). Limestone outcrops at Ao Luek Tai, Ao Luek District, Krabi (8 ° 24 ’ 09.9 ” N, 98 ° 44 ’ 16.3 ” E): CUMZ 7221 (four specimens in ethanol). Limestone outcrops near Than Bok Khorani, Ao Luek District, Krabi (8 ° 22 ’ 28.7 ” N 98 ° 44 ’ 07.9 ” E): CUMZ 7222 (one shell), CUMZ 7223 (10 shells and four specimens in ethanol), and CUMZ 7224 (five specimens in ethanol). Tham Nalakiring Bureau of Monks, Plai Phraya District, Krabi (8 ° 33 ’ 35.3 ” N 98 ° 51 ’ 37.5 ” E): CUMZ 7228 (two specimens in ethanol). Wat Khao Phra, Phrasaeng District, Surat Thani (8 ° 37 ’ 33.0 ” N 98 ° 56 ’ 53.5 ” E): CUMZ 7229 (one specimen in ethanol). Figure 15 Genitalia and spermatophore of Taphrenalla alba sp. nov. specimen CUMZ 7223. A. whole reproductive system. B. internal wall of penis with penial verge. C. spermatophore; inset showing two spines located close to the sperm sac. Type locality. Wat Tham Seua (Tiger Cave Temple), Mueang District, Krabi, Thailand (8 ° 07 ’ 32.9 ” N, 98 ° 55 ’ 27.1 ” E).	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	etymology	Etymology. The specific epithet ‘ alba ’ is from the Greek word ‘ albus ’ meaning ‘ white, whitish’, and refers to the characteristics of the shell having a white line or white subsutural band below the suture.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	diagnosis	Diagnosis. Suture shallowly impressed and with white subsutural band. Epiphallic caecum short and bent at tip. Penial caecum short. Inner sculpture of proximal penis with very soft longitudinal folds then transition to small cuboidal pilasters distally. Dart apparatus located at vagina and penis junction. Spermatophore with two spines on tail filament close to sperm.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	description	Description Shell (fig. 13 C). Shell globosely depressed, medium-sized (width up to 18.5 mm; height up to 10.6 mm) and reddish-brown. Whorls 5 – 6; suture shallowly impressed and with narrow whitish subsutural band. Upper shell surface smooth and without radial grooves. Spire conic; apex raised with a spire angle of about 126 – 144 °. Last whorl well-rounded at periphery. Aperture lip simple; umbilicus narrowly opened. Genital organs (fig. 15). Atrium short. Penis cylindrical and nearly one and a-half times total vagina length; penial caecum present. Internal sculpture of proximal penis with thin longitudinal folds then transformed to small cuboidal penial pilasters in middle and surrounding penial verge in distal penis. Epiphallus slender tube, slightly longer than total penis length. Epiphallic caecum short and bent at the tip, and penial retractor muscle connected near tip. Flagellum slightly long (about half-length of epiphallus) with blunt tip. Vas deferens small and tube-shaped. Vagina prolonged and cylindrical-shaped. Dart apparatus located at junction between vagina and penis. Gametolytic sac elongate bulbous; gametolytic duct long and tube-shaped. Spermatophore elongate and needle-shaped. Tail filament very long tube; region close to sperm sac bearing two spines; furthest spine from sperm sac large and longer than the other. External features (fig. 3). Animal very dark gray in body colour with three milky yellow stripes running from head to caudal horn. One stripe at middle of the body and the other two above foot sole.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
426CF52FA4534B73FD3E84326737BD98.taxon	discussion	Remarks. Taphrenalla alba sp. nov. is similar to T. zemia sp. nov., but can be distinguished by having a reddish-brown shell with narrow whitish subsutural band, whereas T. zemia sp. nov. has a yellowish-brown shell and without subsutural band. Furthermore, the dart apparatus of T. alba sp. nov. is located at the vagina and penis junction, whereas in T. zemia sp. nov. it is located on vagina away from the vagina and penis junction.	en	Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros, Jeratthitikul, Ekgachai, Panha, Somsak (2021): Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21-69, DOI: 10.1163/18759866-BJA10013, URL: http://dx.doi.org/10.1163/18759866-bja10013
