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Nahok, Benchawan, Tumpeesuwan, Chanidaporn, Tumpeesuwan, Sakboworn (2021): Two new species of genus Anceyoconcha S. Tumpeesuwan & C. Tumpeesuwan, in Nahok et al., 2020 (Gastropoda: Pulmonata: Camaenidae), from northeastern Thailand. Raffles Bulletin of Zoology 69: 555-569, DOI: 10.26107/RBZ-2021-0072

Landouria tuberculata sp. nov. urn:lsid:zoobank.org:act: 84133944-D5C9-49A3-9997-8E1C7B335938 Figs 3B, 4B, 5B, 6D–F; Tables 1–4 Diagnosis Shell moderately keeled, with numerous tiny tubercles on shell surface (Fig. 4B). Flagellum like a short protrusion with a pointed tip, penis swollen, base of gametolytic sac narrow. Radula with slender, lanceolate central teeth. Etymology The specific epithet ‘ tuberculata ’ refers to the shell sculpture consisting of numerous tiny tubercles (Fig. 4B). Type material Holotype THAILAND • shell (SH = 9.6 mm, SW = 14.4 mm, AH = 5.2 mm, AW = 5.5 mm, UW = 3.5 mm); Loei Province, Phu Rua District, Wat Thepnimit; 17°26′15.09″ N, 101°16′41.46″ E; alt. 647 m; 23 Sep. 2017; B. Nahok, K. Tanmuangpak and U. Chanlabut leg.; NHMSU-00025. (Fig. 3B) Paratypes THAILAND • 10 shells, 7 living specs; same collection data as for holotype; GenBank No. MN449402 and MZ435747; NHMSU-00026. Description SHELL (Fig. 3B, Table 1). Dextral, medium-sized, tightly coiled, with 6½ slightly convex whorls. Suture rather shallow, apex obtuse with high conical spire. Body whorl angular, with tubercles all over its surface, without distinct incised spiral lines. Colour red brown in living specimens (Fig. 4B), empty shells are paler (Fig. 3B). Umbilicus deep and wide. Aperture obliquely oval, reflexed, and slightly thickened. GENITAL SYSTEM (n = 3) (Fig. 5B, Table 4). Penis cylindrical, swollen in the middle part, internally with rather wavy, corrugated longitudinal pilasters and a short, hollow verge with 2–3 lobes around its opening. Epiphallus evenly cylindrical, as long as penis. Flagellum like a short protrusion with a pointed tip, without nodes, internally with longitudinal slender pilasters. Vas deferens cylindrical, narrow, entering in the lower half of the flagellum. Vagina relatively short, internally with five corrugated longitudinal pilasters. Free oviduct very short. Gametolytic sac slightly thicker at base, with a long, narrow and thin cylindrical tube, and at its distal end a small, swollen spherical sac. Prostate gland very long. Uterus long and thin. RADULA (n = 3). Comprises 126–130 transverse rows with 79–87 teeth per row; radular formula: (28– 30)-(11–13)-1-(11–13)-(28–30). Central tooth symmetric, slender lanceolate, and unicuspid (Fig. 6D). Lateral teeth larger than the central tooth, lanceolate, bicuspid, with large endocone and sharp cusp (Fig. 6D). Marginal teeth tricuspid, with small sharp endocone; large, slender mesocone; and small v-shaped ectocone with two cusps (Fig. 6E–F). Remarks Landouria tuberculata sp. nov. differs from other Thai species of Landouria, by the shell sculpture of numerous tiny tubercles (Fig. 4B). Lateral teeth of radula bicuspid and lanceolate (Fig. 6D–E). Distribution Landouria tuberculata sp. nov. is currently only known from the area surrounding Phu Rua National Park, Phu Rua District, Loei Province, Thailand.

Landouria chloritoides sp. nov. urn:lsid:zoobank.org:act: 740CA614-D17B-47A3-A719-7DFC7D2CE2D5 Figs 3D, 4D, 5D, 6J–L; Tables 1–4 Landouria sp.1 – Ounchareon 2015: 29–32, 44–50, 54–56, 59, 61, figs 4.2c, 4.3c, 4.4. 4.5i–l, 5.5, 5.7. Landouria sp.2 – Buddharaksa 2016: 37–38, 41–42, figs 4.19, 4.20, 5.1, 5.2. Landouria sp.4 – Yingkhamhang 2016: 25–26, 39–41, 43–45, 49–52, figs 4.7, 4.8, 4.21d, 4.22d, 5.3a, 5.4a. Diagnosis Shell shape similar to that of the genus Chloritis Beck, 1837 (Camaenidae), but with last whorl stout and usually slightly angular, peripheral keel blunt, suture deep, aperture oval and oblique. Flagellum small ovate, penis very large and stout compared to vagina; vagina and free oviduct short (Fig. 5D). Radula with lanceolate teeth. Etymology The specific epithet ‘ chloritoides ’ refers to the conchological similarity between the shell of the new species and that of the camaenid genus Chloritis. Type material Holotype THAILAND • shell (SH = 12.0 mm, SW = 19.0 mm, AH = 7.7 mm, AW = 7.3 mm, UW = 3.9 mm); Khon Kaen Province, Chum Phae District, Wat Thep Udom Wanaram; 16°45′29″ N, 101°57′55.03″ E; alt. 406 m; 9 Jun. 2017; B. Nahok, S. Tumpeesuwan and C. Tumpeesuwan leg.; NHMSU-00029. (Fig. 3D) Paratypes THAILAND • 8 shells, 14 living specs; same collection data as for holotype; GenBank No. MN449404 and MZ435749; NHMSU-00030. Other material examined THAILAND • 4 shells, 8 living specs; North East, Khon Kaen Province, Chum Phae District, Pu Lup Cave; 16°49′49.78″ N, 101°59′9.97″ E; alt. 336 m; NHMSU-00031. Description SHELL (Fig. 3D, Table 1). Dextral, somewhat depressed, relatively large-sized. Whorls 6½, suture deep, apex obtuse, and moderately depressed, with low conical spire. Body whorl rounded, slightly keeled, red-brown in living specimens (Fig. 4D), empty shells are paler (Fig. 3D). Umbilicus very deep and wide. Aperture almost circular, with slightly thickened and weakly expanded lip. GENITAL SYSTEM (n = 3) (Fig. 5D, Table 4). Penis very large and stout, longer than vagina, cylindrical, gradually tapering towards epiphallus, internally with six rather thick, corrugated longitudinal pilasters; opening of the short, grooved verge surrounded by 2–3 lobes. Epiphallus short and thick. Flagellum short, elongate ovate, without nodes, internally with three large longitudinal pilasters. Vas deferens long and slender, entering in the lower half of the flagellum. Vagina short, as long as free oviduct, internally with seven, corrugated longitudinal pilasteres varying in size. Gametolytic sac thickened at base, with a long narrow cylindrical tube, and a small oval sac at distal end. Uterus swollen, prostate gland long. RADULA (n = 3). Comprises 125–129 transverse rows with 75–83 teeth per row; radular formula: (26–28)- (11–13)-1-(11–13)-(26–28). Central tooth short lanceolate, symmetric, unicuspid (Fig. 6J). Lateral teeth large and elongated lanceolate (Fig. 6J). Marginal teeth lanceolate, gradually changing from unicuspid to bicuspid and finally tricuspid; endocone small; ectocone slightly oblique, with wide triangular notch and two to four lateral cusps (Fig. 6K–L). Remarks Landouria chloritoides sp. nov. can be distinguished from other species of Landouria in Thailand by its conchological similarity to the genus Chloritis and its much larger and stout penis (Fig. 5D). The radula of L. chloritoides sp. nov. is similar to that of L. tuberculata sp. nov. but differs by its unicuspid lateral teeth (Fig. 5J) vs the bicuspid lateral teeth of L. tuberculata sp. nov. Distribution Landouria chloritoides sp. nov. is currently known only from the limestone area around Phu Pha Man National Park, Chum Phae District, Khon Kaen Province, Thailand.

Key to species by shell characters of Landouria species in northeastern Thailand Finally, in order to facilitate future field research, we provide a dichotomous key for the identification of the currently known Landouria species in northeast Thailand. 1. Shell lustrous, peripheral keel absent, brown band present............................................................................................................................................... L. diplogramma (Möllendorff, 1902) comb. nov. – Shell not lustrous, peripheral keel absent, brown band absent......................................................... 2 2. Peripheral keel blunt...................................................................................... L. chloritoides sp. nov. – Peripheral keel sharp......................................................................................................................... 3 3. Peripheral keel slightly sharp, shell surface with numerous tiny tubercles.... L. tuberculata sp. nov. – Peripheral keel very sharp, shell surface without tubercles.............................................................. 4 4. Shell relatively large with low spire, shell surface with radial scaly processes............................................................................................................................................... L. trochomorphoides sp. nov. – Shell relatively moderate in size with high spire, shell surface without radial scaly processes....... 5 5. Keel more sharp and distinct with its downward bent rim................................... L. elegans sp. nov. – Keel sharp, without downward bent rim........................................................................................... 6 6. Suture indented, growth line obvious................................................................................................................................................................ L. strobiloides C. Tumpeesuwan & S. Tumpeesuwan, 2019 – Suture not indented, growth line obscure........................................................... L. circinata sp. nov., Published as part of Nahok, Benchawan, Tumpeesuwan, Sakboworn & Tumpeesuwan, Chanidaporn, 2021, Five new species of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from northeastern Thailand, with note on genitalia and radula morphology of Landouria diplogramma (Möllendorff, 1902), pp. 142-166 in European Journal of Taxonomy 767 (1) on page 162, DOI: 10.5852/ejt.2021.767.1495, http://zenodo.org/record/5514839, {"references":["Mollendorff O. F. 1902. Neue Arten und Unterarten von Fruhstorfer in Siam gesammelt. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 34 (9 - 10): 153 - 160.","Tumpeesuwan C. & Tumpeesuwan S. 2019. First verified record of the genus Landouria Godwin- Austen, 1918 from Thailand (Gastropoda: Stylommatophora: Camaenidae) with description of a new species. Raffles Bulletin of Zoology 67: 298 - 305. https: // doi. org / 10.26107 / RBZ- 2019 - 0021"]}

Landouria diplogramma (M��llendorff, 1902) comb. nov. Figs 3F, 4F, 5F, 6P���R; Tables 1���4 Plectotropis diplogramme M��llendorff, 1902: 158���159. Ganesella diplogramme ��� Panha 1996: 36. Aegista (Plectotropis) diplogramma ��� Hemmen & Hemmen 2001: 47. Thaitropis sp. ��� Sutcharit et al. 2017: 249, fig. 6���32e. Diagnosis Shell small, low-conical, weakly keeled, with a slightly elevated spire, whitish corneous with a pale brown band above periphery. Flagellum like a short protrusion, with rounded tip. Penis short swollen. Central and the first few lateral radular teeth triangular. Material examined THAILAND ��� 29 shells, 14 living specs; Nakhon Ratchasima Province, Pak Chong District, Wat Thep Phithak Punnaram; 14��36���54.57��� N, 101��15���55.25��� E; alt. 451 m; 23 Jun. 2017; B. Nahok and U. Chanlabut leg.; GenBank No. MN449411 and MZ435752; NHMSU-00034. (Fig. 3F) Description SHELL (Fig. 3F, Table 1). Dextral, small, depressed conical. Whorls 5��, suture rather shallow, apex obtuse and depressed, whitish corneous, with a pale brown peripheral band. Protoconch almost smooth. Umbilicus moderately deep and wide. Aperture oval oblique, with thin, solid, and weakly reflected lip. GENITAL SYSTEM (n = 3) (Fig. 5F, Table 4). Penis divided into two short portions, the portion that connects to epiphallus is swollen and the portion that connects to atrium is constricted; internally with three corrugated longitudinal pilasters; with short, rather rounded verge. Flagellum like a short protrusion, internally with two narrow longitudinal pilasters. Vas deferens long, cylindrical, entering the epiphallus at base. Free oviduct very short. Vagina twice as long as free oviduct, internally with five thick, corrugated longitudinal pilasters. Proximal part of gametolytic sac slightly thicker than the cylindrical tube, with a medium-sized oval sax at distal end. Uterus and prostate gland long. RADULA (n = 3). Comprises 121���125 transverse rows with 61���69 teeth per row; radular formula: (23���25)- (7���9)-1-(7���9)-(23���25). Central tooth usually symmetric, unicuspid, triangular (Fig. 6P). Lateral teeth longer and larger than central teeth, bicuspid, ectocone small, with sharp cusp (Fig. 6Q). Marginal teeth gradually changing from bicuspid to tricuspid, small endocone; mesocone large lanceolate; ectocone triangular with two to four tiny cusps (Fig. 6R). Remarks Specimens deposited in the Leiden Museum, The Netherlands, RMNH.MOL.309867 and RMNH. MOL.309851 (https://images.app.goo.gl/PgrpqKaWuSrgjdvN9), are identified as Plectotropis diplogramme M��llendorff, 1902. These specimens appear identical to specimens from Khao See Siad Ah in Nakhon Ratchasima Province, therefore we used the name ��� Landouria diplogramma (M��llendorff, 1902) ��� for this species in the present study. Landouria diplogramma differs from other Thai species of Landouria by its relatively small lustrous shell without peripheral keel, but with a pale brown band present above periphery (Figs 3F, 4F). Its radular teeth are relatively short (Fig. 6P���R). Its penis and flagellum are relatively short (Fig. 5F). The genus Thaitropis Schileyko, 2004 was recently reported in Thailand (Schileyko 2004; Sutcharit et al. 2017). Yet, this genus was synonymized with Landouria by Nurinsiyah et al. (2019). Based on the illustration provided by Sutcharit et al. (2017: fig. 6���32e), we identify their Thai Thaitropis sp. as L. diplogramma, because it has the same shell morphology and colours: Plectotropis diplogramme M��llendorff, 1902, RMNH.MOL.309867, and L. diplogramma (M��llendorff, 1902) from Khao See Siad Ah, Nakhon Ratchasima (NHMSU-00034) (Figs 3F, 4F). Unfortunately, the whereabouts of the type specimen of Plectotropis diplogramme are unknown. Yet, it might be deposited in the collections of the Senckenberg Museum in Frankfurt, Germany where it should be looked up and checked in a further study. Distribution Landouria diplogramma was collected by local people in the mountain range known as Khao See Siad Ah, Pak Chong District, Nakhon Ratchasima Province, Thailand. Other reported localities comprise Bangkok, Hinlap, Muoklek, Kanburi (Kanchanaburi) (M��llendorff 1902)., Published as part of Nahok, Benchawan, Tumpeesuwan, Sakboworn & Tumpeesuwan, Chanidaporn, 2021, Five new species of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from northeastern Thailand, with note on genitalia and radula morphology of Landouria diplogramma (M��llendorff, 1902), pp. 142-166 in European Journal of Taxonomy 767 (1) on pages 159-161, DOI: 10.5852/ejt.2021.767.1495, http://zenodo.org/record/5514839, {"references":["Mollendorff O. F. 1902. Neue Arten und Unterarten von Fruhstorfer in Siam gesammelt. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 34 (9 - 10): 153 - 160.","Panha S. 1996 A checklist and classification of the terrestrial pulmonate snails of Thailand. Walkerana 8 (19): 31 - 40.","Hemmen J. & Hemmen C. 2001 Aktualisierte Liste der terrestrischen Gastropoden Thailands. Schriften zur Malakozoologie 18: 35 - 70.","Sutcharit C., Tongkerd P. & Panha S. 2017. Land Snails the Invaluable Bio-Resources for the Kingdom of Thailand. The Thailand Research Fund, Bangkok. [In Thai.]","Schileyko A. A. 2004. Treatise on recent terrestrial pulmonate molluscs. Part 12. Bradybaenidae, Monadeniidae, Xanthonychidae, Epiphragmophoridae, Helminthoglyptidae, Elonidae, Humboldtianidae, Sphincterochilidae, Cochlicellidae. Ruthenica Supplement 2: 1627 - 1764.","Nurinsiyah A. S., Neiber M. T. & Hausdorf B. 2019. Revision of the land snail genus Landouria Godwin- Austen, 1918 (Gastropoda, Camaenidae) from Java. European Journal of Taxonomy 526: 1 - 73. https: // doi. org / 10.5852 / ejt. 2019.526"]}

Landouria elegans sp. nov. urn:lsid:zoobank.org:act: 6A33A2BA-7197-4C1E-AEB3-D7815F8A05AC Figs 3E, 4E, 5E, 6M���O; Tables 1���4 Landouria sp.3 ��� Yingkhamhang 2016: 23���24, 39���45, 47���48, 61, 74, figs 4.5, 4.6, 4.21���4.23c, 5.1c, 5.2c. Diagnosis Shell similar to Landouria circinata sp. nov., but with a sharper and more prominent keel at the periphery and dark brown zones on the periphery, just above and below the keel. Flagellum droplet shaped, penis long and cylindrical, vagina and free oviduct very short. Radula with elongate tongue-shaped central and lateral teeth. Etymology The specific epithet ��� elegans ��� (Latin for ���magnificent���) refers to the beauty of the shell. Type material Holotype THAILAND ��� shell (SH 10.0 = mm, SW 17.7 = mm, AH = 6.4 mm, AW = 7.1 mm, UW = 3.6 mm); Khon Kaen Province, Phu Pha Man District, Phu (mountain) Hua Chang; 16��39���10.54��� N, 101��48���6.67��� E; alt. 305 m; 3 May 2018; B. Nahok, S. Tumpeesuwan and C. Tumpeesuwan leg.; NHMSU-00032. (Fig. 3E) Paratypes THAILAND ��� 11 shells; 9 living specs; same collection data as for holotype; GenBank No. MN449408 and MZ435751; NHMSU-00033. Description SHELL (Fig. 3E, Table1). Dextral, conical, relatively large-sized.Whorls 6. Protoconch with radially elongated wrinkles. Suture rather shallow, apex obtuse depressed, with rather high conical spire. Last whorl with a sharp keel that is distinctly bent downwards. Colour brownish-corneous with a dark brown zone bordering the keel at the periphery. Umbilicus very deep and wide. Apertural lip thickened, rounded rhombic. GENITAL SYSTEM (n = 3) (Fig. 5E, Table 4). Penis longer than vagina, cylindrical, internally with four corrugated, longitudinal pilasters; verge absent. Epiphallus shorter than penis, distal part (ep2) small, whereas proximal part (ep1) is swollen. Flagellum droplet-shaped, without nodes, internally with four corrugated, longitudinal pilasters. Vas deferens long, cylindrical, proximal part that connects to free oviduct somewhat thicker than distal part, entering the flagellum at its base. Vagina very short, internally with five corrugated longitudinal pilasters. Free oviduct shorter than vagina. Proximal part of gametolytic sac rather stout, but tapering to become a slender tube with large oval sac at distal end. Uterus and prostate gland very long and large. RADULA (n = 3). Comprises 110���114 transverse rows with 69���77 teeth per row; radular formula: (22���24)- (12���14)-1-(12���14)-(22���24). Central and lateral teeth unicuspid, tongue-shaped. Central tooth smaller and shorter than lateral teeth (Fig. 6M���N). Marginal teeth asymmetric, tricuspid with small endocone; mesocone large with curved margins; ectocone triangular and located at tooth base (Fig. 6N���O). Remarks Landouria elegans sp. nov. is conchologically quite similar to L. circinata sp. nov. and L. strobiloides, from which it differs by its sharper and more distinct keel with its downward bent rim (Fig. 3A vs Fig. 3E). The radula of L. elegans sp. nov. resembles that of L. circinata sp. nov., but central tooth and lateral teeth of L. elegans sp. nov. are longer and more slender than in L. circinata sp. nov. (Fig. 6A���C vs Fig. 6M���O). Epiphallus of L. elegans sp. nov. is similar to that of L. chloritoides sp. nov., but in L. elegans sp. nov. it is larger and connects with the vas deferens near the flagellum base (Fig. 5D vs Fig. 5E). Distribution Landouria elegans sp. nov. is known only from the type locality., Published as part of Nahok, Benchawan, Tumpeesuwan, Sakboworn & Tumpeesuwan, Chanidaporn, 2021, Five new species of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from northeastern Thailand, with note on genitalia and radula morphology of Landouria diplogramma (M��llendorff, 1902), pp. 142-166 in European Journal of Taxonomy 767 (1) on pages 158-159, DOI: 10.5852/ejt.2021.767.1495, http://zenodo.org/record/5514839, {"references":["Yingkhamhang A. 2016. Shell Morphology and Reproductive Anatomy of Land Snails Genus Landouria Godwin-Austen, 1918 in Northeastern Thailand. Unpublished Bachelor's Degree Science Project. Department Faculty of Science, Mahasakham University, Maha Sarakham. [In Thai with English abstract.]"]}