Search

Pravonitor insularum (Iredale, 1937) (Figures 4, 14 (a)) Nitor (Pravonitor) kreffti insularum Iredale, 1937: 3. Material examined Holotype. AM C.29647, Mer Is., Torres Strait, Far North Qld, Australia. Paratypes. AM C. 170795 156 specimens; AM C.104171 1 specimen. Other material. See Table 1. Diagnosis External morphology. Shell (Figure 14 (a)) medium-sized (SW 11.3–13.6 mm), pale golden with a pale orange-brown supraperipheral band, 5.2–5.5 whorls, discoidal with a low conical spire, whorls rounded; umbilicus very narrowly open. Protoconch and teleoconch sculptured with very fine spiral grooves. Remarks The first name to be applied to Torres Strait Pravonitor species was Helix (Thalassia) annulus Brazier, 1876 from New Guinea (Brazier 1876). This name was restricted to New Guinean populations by Iredale (1937), and the new subspecies Nitor (Pravonitor) kreffti insularum was introduced for Torres Strait Island populations (type locality Mer Island). The new subspecies was originally described as differing from P. krefftii (type locality Cape York) in its slightly smaller size, more depressed spire and the presence of a coloured anteperipheral band (Iredale 1937). The two subspecies were synonymised by Smith (1992) based on the original descriptions and an examination of types. This was upheld by Hyman and Ponder (2010), who stated that shell size and the presence of a coloured band is highly variable and found no clear difference between the shells of mainland and island forms, and illustrated specimens from Gerbar Island under the name Pravonitor kreffti. Stanisic (2008) recorded P. kreffti from Mua, Badu, Iama, Erub, Mer and Waibene Islands as well as Cape York, based on the collections of the Australian Museum and Queensland Museum. However, in the current study we found that specimens from Iama Island and Gerbar Island (situated midway between Australia and New Guinea) differed considerably in their genital morphology and mitochondrial DNA to specimens from Mua, Badu, Waiben and Muralug Islands, which are much closer to Australia. Specimens from Cape York matched the latter group. Comparison of type material (where present) and original descriptions confirmed that these two species corresponded to Pravonitor annulus and Pravonitor kreffti. The available material from Mer Island was too old for successful DNA extraction and sequencing, and all preserved specimens were subadult. Dissection demonstrated that the specimens belong to the genus Pravonitor, but did not provide sufficient information for species assignment. Shell measurements showed that these shells were significantly flatter than those of all other Torres Strait Island groups, as suggested by Iredale (1937). However, the available shells were also slightly lower in whorl count than those of other island populations (see Table 4), possibly inflating the perceived difference, since in land snails shell growth is allometric and relative height generally increases with whorl count (Goodfriend 1986). Examination of the shells also confirmed the presence of an anteperipheral band, also present in Pravonitor annulus but not P. kreffti. In the absence of anatomical and molecular data, the only evidence that P. insularum is distinct from the other Torres Strait species lies in the significantly flatter shell and the presence of an anteperipheral band. There is considerable evidence to show that shell shape is often related to habitat preference rather than shared ancestry (e.g. Köhler et al. 2008; Stankowski 2011; Criscione and Köhler 2013). However, Mer Island is also more distant than the other islands, and has a different origin, having been formed from volcanic activity rather than being part of the continental shelf. Given that different species exist on the mainland/Inner islands/Near Western islands and on the Central islands/Top Western islands/New Guinea, it appears mostly likely that the isolated Mer Island also harbours a unique species. Pravonitor insularum is currently placed in synonymy with P. kreffti, but is more similar to and lies geographically closer to P. annulus. Herein we remove P. insularum from synonymy with P. kreffti; however, we do not feel there is sufficient evidence to place it in synonymy with P. annulus, and therefore we retain this name for the Mer Island populations alone. The Eastern group of islands also contains Dauar Island, Waier Island, Erub Island and Ugar (Stephens) Island. Four shells from Erub Island were measured and were found to differ significantly from the Mer Island shells in both size and shape (SW, SH, SW/SH: p P. annulus or P. kreffti. It is possible that the Eastern Islands may contain more than one species., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1773-1775, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Brazier J. 1876. List of land shells collected during the Chevert expedition. Proc Linn Soc NSW. 1: 117 - 133. doi: 10.5962 / bhl. part. 12392","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1.","Stanisic J. 2008. The land snails of Mua Island. Mem Queensl Mus, Cult Herit Ser. 4 (2): 491 - 494.","Goodfriend GA. 1986. Variation in land-snail shell form and size and its causes: a review. Syst Biol. 35 (2): 204 - 223. doi: 10.1093 / sysbio / 35.2.204.","Kohler F, Brinkmann N, Glaubrecht M. 2008. Convergence caused confusion: on the systematics of the freshwater gastropod Sulcospira pisum (Brot, 1868) (Cerithioidea, Pachychilidae). Malacologia. 50 (1): 331 - 339. doi: 10.4002 / 0076 - 2997 - 50.1 - 2.331.","Stankowski S. 2011. Extreme, continuous variation in an island snail: local diversification and association of shell form with the current environment. Biol J Linn Soc. 104 (4): 756 - 769. doi: 10. 1111 / j. 1095 - 8312.2011.01748. x.","Criscione F, Kohler F. 2013. Conserved shell disguises diversity in Mesodontrachia land snails from the Australian Monsoon Tropics (Gastropoda: Camaenidae). Zool Scr. 42 (4): 389 - 405. doi: 10.1111 / zsc. 12011."]}

Pravonitor stuarti sp. nov. (Figures 4, 14 (i), 21) Etymology Named for Stuart Palethorpe. Material examined Holotype. QM MO86002, High Range, Far North Qld (13.05° S, 143.45° E), 3 November 2012, K. Aland. Paratype. QM MO80213 (same data as holotype). Other material examined. See Table 1. Diagnosis External morphology. Shell (Figure 14 (i)) medium-sized (SW 12.1–13.6 mm), pale gold above, pale amber below, with an amber supraperipheral band, 5.2–5.5 whorls, discoidal with a moderately elevated, slightly concave spire, whorls rounded, umbilicus very narrow, sealed with a mucous plug, partially covered by reflected lip. Protoconch and teleoconch sculptured with very fine spiral grooves, hardly visible on protoconch. Body cream, with dark grey tail (incl. caudal horn), pale grey head, pale sole, grey band along side of foot (in alcohol). Right and median mantle lobes moderate, left lobe moderately small; right lobe grey and cream, median lobe dark grey, left lobe cream; shell lappets long, wide at base, tapering to a point. Caudal horn very long. Genital anatomy. Genitalia (Figure 21) with short vagina; bursa copulatrix moderately long, duct broad at base, becoming narrower, coiled, bursa elongate, internally sculptured with transverse ridges basally and longitudinal ridges in duct. Penis very long and narrow, tubular, slightly broader at base, narrower in middle, broadening slightly after exiting penial tunica and slightly swollen near epiphallic boundary; internally with multiple narrow longitudinal pilasters; approx. 80% enclosed in penial tunica. Two penial caecae inserted on penis near base, inside penial tunica; one short, one longer. Epiphallus enters penis through a small verge; epiphallus slightly longer than penis; epiphallus 2 longer than epiphallus 1; long epiphallic caecum present, with medial attachment of the penial retractor muscle; flagellum long, slender, slightly swollen terminally, with no obvious internal cryptae. Remarks Pravonitor stuarti is very similar in appearance to P. ferrugineus, and was originally identified as belonging to this species. However, in the current study a population of P. ferrugineus from High Range was found to have a larger and flatter shell with a more concave spire, while still exhibiting the typical cream-coloured shell with a reddish brown supraperipheral band characteristic of the species. Dissection showed that these specimens differed in penis size, shape and interior sculpture, and in the shape of the flagellum. In particular, while the penis was still long and slender with a thick penial tunica, it was longer and more uniformly narrow in P. stuarti, with a small penial verge and interior sculpture of longitudinal ridges rather than the pointed pustules seen in P. ferrugineus. In addition, the flagellum of P. stuarti is slender and uncoiled, while that of P. ferrugineus is coiled and shows some signs of small internal cryptae, indicating the presence of a coiled spermatophore with small spines (similar to that seen in P. annulus). A single specimen of P. stuarti has been sequenced, and shows it to be the basal member of genus Pravonitor, but also that it is nearly identical to P. ferrugineus. Despite this, we feel that the very distinct reproductive system, particularly the strikingly distinct sculpture of the inner penial wall, indicates that the two morphospecies must be reproductively isolated. Only a single spirit-preserved lot of P. stuarti is known, from High Range, south of Lockhart River on Cape York peninsula (Figure 4). Pravonitor ferrugineus is found slightly to the north, in Iron Range National Park. It is possible that the two species are sympatric, but currently very little data is available for either species. The very distinct differences in the penial sculpture seen in the two species would ensure reproductive isolation, even in sympatry. Increased collecting on Cape York is needed to clarify the range of this species. Pravonitor stuarti can be distinguished from most other species by its cream shell and rust-coloured supraperipheral band, and from Pravonitor ferrugineus by its larger size, flatter shell and more concave spire, as well as differences in its genital morphology., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1788-1790, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681

Geminitor kullaensis sp. nov. (Figures 4, 6 (d), 11) Etymology Named for KULLA (McIlwraith Range) National Park, where the species is found. KULLA is an acronym for Kaanju, Umpila, Lama Lama and Ayapathu, referring to the four clan groups who are the traditional owners of the land. Material examined Holotype. QM MO85998, Coen, c. 38 km SSE at Kendle R. Crossing (14.26°S, 143.29°E), Qld, Australia, R. and N. Crookshanks, 26 September 2000. Paratypes. QM MO 68617 (same data as holotype). Other material. See Table 1. Diagnosis External morphology. Shell (Figure 6 (d)) small (SW 10.6 mm), pale gold, 5.9 whorls, subdiscoidal with a moderately elevated spire, whorls rounded; umbilicus narrow, sealed; protoconch with moderately coarse spiral grooves, becoming finer and more crowded on teleoconch. Body white (in alcohol). Mantle lobes small; shell lappets moderately large, triangular, tapering. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 11) with short vagina; bursa copulatrix moderately short, duct broad basally, internally with v-shaped ridges, bursa oval. Free oviduct very long. Penial complex very long (caecum sits behind albumen gland), penis internally with two chambers, similar in length; distal portion sculptured with two high, raised longitudinal pilasters, joined together proximally, tapering distally; proximal portion sculptured with transverse elongate plates arranged like petals around opening to epiphallus; 30% enclosed in penial tunica. Epiphallus enters penis through simple pore; epiphallus equal in length to penis, epiphallus 2 similar in width to penis, gradually narrowing, epiphallus 1 narrow; epiphallus 1 longer than epiphallus 2; long epiphallic caecum present, with basal attachment of the penial retractor muscle; flagellum moderately short, slender, without obvious internal cryptae. Remarks Geminitor kullaensis was preliminarily identified as Helicarionidae CY 13 through curatorial work and is known from south of Coen to the McIlwraith Range in northern Qld (Figure 4). An additional single subadult specimen from 300 km farther south at Bloomfield River was tentatively identified as G. kullaensis. If this identification is correct, the range of G. kullaensis is significantly expanded, being found both north and south of the Normanby Basin. However, the distribution of this species needs to be confirmed with adult material. According to our phylogenetic analysis (which, however, excludes G. macveae), the nearest relative is another new species, G. wenlockensis from farther north at Wenlock River. Geminitor kullaensis can be clearly identified as belonging to the genus Geminitor by its two-chambered penis and its long caecum with basal attachment of the penial retractor muscle. It differs from its congeners in its much longer penial complex, with a longer proximal penial chamber. Geminitor kullaensis can be distinguished from G. villaris, G. macveae and G. wenlockensis by its larger size, and from G. laura by its slightly smaller size and subdiscoidal rather than discoidal shell. v, Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1765-1767, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681

Pravonitor monteithi (Stanisic, 2010) (Figures 4, 14 (g), 19) Malandena monteithi Stanisic, 2010: 296–297. Holotype. QMMO48587, Isley Hills, north-eastern Qld, Australia (17°03ʹS, 145°42ʹE), rainforest, 1050 m, under logs, 30 November 1993, G. Monteith, H. Janetzki, D. Cook. Paratypes. QMMO24798, QMMO48164, QMMO48203, QMMO48693, QMMO78927, AM C.425580. Other material. See Table 1. Diagnosis External morphology. Shell (Figure 14 (g)) medium-sized (SW 11.9–13.5 mm), amber with a darker underside, 5.1–5.4 whorls, subdiscoidal with a low spire, whorl profile rounded; microsculpture of very fine to obsolete spirals on protoconch and teleoconch; umbilicus very narrow, partially covered by reflected lip. Body white with black eyestalks (in alcohol). Mantle lobes moderately small; shell lappets moderately large, narrow, tapering. Caudal horn large. Genital anatomy. Genitalia (Figure 19) with short vagina, internally with longitudinal pilasters; bursa copulatrix moderately short, duct broad at base, then narrowing, internally with distinct longitudinal pilasters; bursa elongate. Penis moderately small, bulging at base, tubular medially, swollen proximally with large blind tip; internally with 1–2 pustulose longitudinal pilasters, internal wall sculptured with fine pustules; basal bulge with thickened, folded region internally; basal bulge not enclosed in penial tunica. Epiphallus enters penis through a simple pore; epiphallus longer than penis; epiphallus 2 longer than epiphallus 1; moderately long epiphallic caecum present, with medial attachment of the penial retractor muscle; flagellum long, slender, without obvious internal cryptae. Spermatophore with soft capsule and smooth, elongate tail-pipe with no sculpture. Remarks Pravonitor monteithi was previously included in Malandena (now Elatonitor) and bears a strong external similarity to Elatonitor suturalis and P. aquilonia. It can be distinguished by its smaller size, finer teleoconch sculpture and rounded whorl periphery. The presence of an epiphallic caecum with a medial attachment of the penial retractor muscle and an internally pustulose penis interior definitively places this species in Pravonitor. Only a single specimen could be sequenced,and this sequence diverged from within the clade P. aquilonia. It is possible that P. monteithi should be recognised as a synonym of P. aquilonia; however, there are distinct morphological differences between the two and we believe there is insufficient information to synonymise the two species. Specimens identified in the current study as Pravonitor monteithi had a penis with a large, swollen blind tip, and a basal bulge containing a thickened, folded region, as well as 1–2 penial pilasters. In contrast, P. aquilonia is significantly larger, and has a penis that is narrowed proximally with no blind tip, no penial pilasters, and no thickened, folded region basally.Hence,the two species are retained as separate based on morphological differences despite the close relationship shown in the mitochondrial tree. It is possible that these species have diverged very recently and that the morphological differences are not yet reflected in the tree. This species is recorded from rainforest in the Atherton Tablelands and adjacent coastal ranges (Stanisic et al. 2010); however, in the current study only specimens from Lamb Range and from the adjacent northern parts of the Atherton Tableland (e.g. Tinaroo, Lake Eacham) could be confirmed as belonging to P. monteithi (Figure 4). Other specimens from the Atherton Tableland preliminarily identified as P. monteithi were found to be either M. suturalis or P. aquilonia. Further collecting throughout this area to confirm the distribution of these three very similar species would be of great benefit., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1784-1786, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press."]}

Pravonitor annulus (Brazier, 1876) (Figures 4, 14 (b), 15) Helix (Thalassia) annulus Brazier, 1876: 100. Pravonitor annulus: Iredale 1941: 65. Pravonitor kreffti: Hyman and Ponder 2010: 64–65, figs 17 M–0, 18 G, 19 H–I, 20 G–H, 21 H– I, 22C (in part). Material examined Syntypes. AM C.171131, C.171132, Mawatta (Marrahata) Village, Binaturi (Katow) River mouth, New Guinea, 09° 08’ 15” S, 142° 57’ 00” E, coll. Brazier, July 1975. Probable syntypes. AM C.55456, Binaturi (Katow) River, New Guinea, 09° 08’ 15” S, 142° 57’ 00” E, coll. J. Brazier, July 1875), C. 104171 (Torres Strait, Dungeness Island, 09° 51’ 00” S, 142° 54’ 00” E, coll. J. Brazier, 20 July 1875. Other material. See Table 1. Diagnosis External morphology. Shell (Figure 14 (b)) medium-sized (SW 12.6–15.8 mm), golden brown with a darker supraperipheral band, 5.7–6.2 whorls, subdiscoidal with a subangulate whorl profile; shell microsculpture with fine grooves on the protoconch and teleoconch, becoming obsolete on the teleoconch; umbilicus narrow, sealed with a mucous plug, partially covered by reflected lip. Body cream (in alcohol). Mantle lobes moderately small; shell lappets moderately large, narrow, tapering. Caudal horn large. Genital anatomy. Genitalia (Figure 15) with short vagina, internally with wavy longitudinal pilasters; bursa copulatrix moderately short, duct narrow, internally with wavy transverse pilasters, bursa elongate. Penis tubular, with one longitudinal pilaster, internal wall sculptured with pustules, smaller and rounded proximally, larger and more diamondshaped distally; approx. 75% enclosed in penial tunica. Epiphallus enters penis through a small verge; epiphallus longer than penis; epiphallus 2 longer than epiphallus 1; long epiphallic caecum present, with medial attachment of the penial retractor muscle; flagellum long, spiralling, containing internal cryptae. Remarks This species (preliminarily identified above as P. kreffti ‘Central Islands’) was first introduced as Helix (Thalassia) annulus (Brazier, 1876) for a species from Mawatta Village, on the banks of the Binaturi River, New Guinea (given as Marrahata Village, Katow River), and Giaka (= Dungeness) Island, from the Central group of islands in the Torres Strait. The specimens were collected in 1875 on the Chevert Expedition to New Guinea. Iredale (1937) introduced a replacement name for the Torres Strait Islands populations (Helix (Pravonitor) kreffti insularum, type locality Mer Island), presumably assuming that the New Guinean annulus would not also extend to the Torres Strait Islands. The type material was assumed to be lost and so this species was regarded as not applying to any Australian species (Smith 1992). However, after careful examination of the original material in the current study and comparison with shells from multiple other islands in the Torres Strait, we have concluded that this species is a member of the genus Pravonitor and that it is found at the type locality in New Guinea, as well as on the Top Western and Central groups of Torres Strait Islands, including Dauan, Iama, Gerber and Giaka Islands (Figure 4). The syntypes are mostly juvenile shells, with only three approaching adult size, at 5.1– 5.4 whorls. All specimens have golden, shining shells with a distinct band, and a whorl profile that is rounded above and below a slightly angulate periphery. In the case of both P. insularum and P. annulus, the lack of fresh, adult material from the type localities and from throughout the Torres Strait Islands significantly impedes the process of name clarification. It is possible that the Mer Island specimens may prove to be conspecific with the Central Islands species, in which case P. insularum will become a junior synonym of P. annulus. Our intent is to take the most conservative path and to avoid either introducing new names or placing names in synonymy based on incomplete data. For that reason, we retain P. insularum for Mer Island populations and P. annulus for both New Guinean and Top Western/Central Island populations at the present time, pending further investigation. Pravonitor annulus can be distinguished from P. kreffti (found on the Inner and Near Western Islands) by its slightly smaller and more depressed shell with a subangulate whorl profile and a supraperipheral band, and by differences in the genital anatomy, including the presence of a cylindrical penis with no blind tip, a very long epiphallic caecum with medial attachment of the penis retractor muscle, and a coiled flagellum with internal cryptae. This contrasts to the broad penis with a large blind tip, relatively short epiphallus with the penis retractor muscle attaching at or near the tip, and the straight flagellum with no externally visible cryptae of P. kreffti. It is not clear whether the two species co-exist on any islands; targeted collecting in this region would help to delimit species distributions and clarify the differences between the two species., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1775-1777, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Brazier J. 1876. List of land shells collected during the Chevert expedition. Proc Linn Soc NSW. 1: 117 - 133. doi: 10.5962 / bhl. part. 12392","Iredale T. 1941. Guide to the land shells of New South Wales. Part III. Aust Nat. 11: 1 - 8.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1.","Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408."]}

Geminitor macveae sp. nov. (Figures 4, 6 (e), 12) Etymology Named for Tracey McVea, in acknowledgement of her tireless efforts as Grants and Research Support Officer at the Australian Museum Research Institute. Material examined Holotype. QM MO85999, Rifle Ck, c. 1 km NW of Mt Molloy, north-eastern Qld, Australia (15 57’ S, 145 19’ E), 10 September 1988, J. Stanisic, D. Potter. Paratype. QM MO20694 (same data as holotype). Other material. See Table 1. Diagnosis External morphology. Shell (Figure 6 (e)) small (SW 6.6 mm), golden amber, 5.2 whorls, subdiscoidal with a low conical spire, whorls rounded; umbilicus narrow; protoconch and teleoconch with fine spiral grooves. Body white (in alcohol). Mantle lobes moderately small; shell lappets long, triangular, tapering. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 12) with short vagina; bursa copulatrix of moderate length, duct broad, internally with fine, longitudinal, anastomosing ridges; bursa elongate. Penis short, slender, internally with two chambers, distal portion sculptured with one main longitudinal pilaster and a second, shorter and narrower pilaster; proximal portion sculptured with longitudinal ridges; penial verge absent;penis 100% enclosed in penial tunica.Epiphallus longer than penis, similar in width to penis; two arms of epiphallus approximately equal in length; long epiphallic caecum present, with basal attachment of the penial retractor muscle; flagellum moderately short, slender, coiled, without obvious internal cryptae. Remarks Geminitor mvceae (preliminarily identified as P. villaris ‘Rifle Creek’) is only known from the holotype and paratype, from Rifle Creek near Mt Molloy (Figure 4). This species was previously subsumed within G. villaris, which it closely resembles. However, G. macveae is significantly smaller than G. villaris, with a much smaller and narrower penis and distinctly different internal penial sculpture. It is the smallest and most southerly distributed of all Geminitor species.

Geminitor gen. nov. Type species Pravonitor laura Stanisic, 2010. Etymology From geminus (Latin = twin-born, twin), referring to the double-chambered penis, and Nitor, a helicarionid snail genus; of masculine gender. Differential diagnosis External appearance. Shell small to medium-sized (SW 6.6–14.6 mm), pale gold to amber, often paler on the underside, glossy, 4.8–6.3 whorls, discoidal to subdiscoidal with a low to moderately elevated spire; whorls rounded; umbilicus narrow, sealed with a mucous plug; lip slightly reflected over umbilicus. Protoconch with moderately coarse spiral grooves, becoming finer and more crowded on teleoconch. Body colour pale brown to white, often with a grey tail tip and black eyestalks. Mantle lobes small; shell lappets moderately long, triangular, wide at base, tapering to a point. Caudal horn moderately large. Genital anatomy. Ovotestis embedded in digestive gland. Carrefour visible; talon usually embedded in albumen gland. Spermoviduct not folded. Free oviduct long, with elongate capsular gland; free oviduct internally smooth. Vagina very short. Bursa copulatrix moderately short, inserted on vagina, base of duct usually swollen, then narrowing; bursa elongate. Penis with two chambers, distal chamber with longitudinal pilasters and ridges, upper chamber sculptured with rounded, transverse plates. Verge-like structure sometimes present between penial chambers. Penis tunica attached by muscle fibres to middle of epiphallus; epiphallus enters penis through simple pore or small verge; epiphallus 2 longer than or shorter than epiphallus 1; epiphallus 2 broader than epiphallus 1; epiphallic caecum moderately long to long, penial retractor muscle attached to base of caecum. Long, slender epiphallic flagellum with axial filament present; no internal cryptae visible. Remarks Geminitor is introduced for small to medium-sized snails with a relatively pale, unbanded, discoidal to subdiscoidal shell and a two-chambered penis. Two species are removed from Pravonitor (G. laura, G. villaris) and three species are described as new (G. kullaensis, G. wenlockensis, G. macveae). Geminitor can be characterised by its small to medium-sized, pale gold to amber, glossy shell, often paler on the underside, with rather coarse microspiral sculpture on the protoconch (becoming finer on the teleoconch). The shell is discoidal to subdiscoidal in shape, with a low to moderately elevated spire and a rounded whorl profile. Anatomically, it is distinguished particularly by its two-chambered penis, often with a beehive-shaped verge leading into the second chamber, and/or a series of overlapping plates around the opening between the two chambers. Other key features include the long, slender reproductive system with visible carrefour, the very short vagina, the broad epiphallus 2 compared to epiphallus 1, the long caecum with basal attachment of the penial retractor muscle, and the long, slender flagellum. This genus is restricted to Cape York Peninsula, being found from just north of Cairns to Bramwell. This area is under-collected and it is likely that there are additional undescribed species., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1757-1758, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press."]}

Pravonitor ferrugineus Stanisic, 2010 (Figures 4, 14 (d), 17) Pravonitor ferruginea [sic!] Stanisic, 2010: 300–301. Material examined Holotype. QM MO12752, East Claudie River, Iron Range, Far North Qld, Australia, monsoon rainforest, 26 July 1981, A. Green. Paratypes. AM C.122987, AM C.432173, AM C.432174. Other material. See Table 1. Diagnosis External morphology. Shell (Figure 14 (d)) medium-sized (SW 10.2–13.6 mm), pale cream to golden brown, with an orange-brown supraperipheral band, 5.2–6.4 whorls, discoidal with a low conical spire, whorls rounded, umbilicus very narrow. Protoconch and teleoconch sculptured with very fine spiral grooves. Body cream (in alcohol). Mantle lobes moderately large; shell lappets moderately long, wide at base, tapering to a point. Caudal horn moderately long. Genital anatomy. Genitalia (Figure 17) with moderately short vagina; bursa copulatrix moderately long, duct slightly broader at base, tapering, bursa not distinct, internally sculptured with longitudinal ridges. Penis moderately long, tubular, swollen just before epiphallic boundary with a small blind tip; internally with long, pointed, conical pustules, becoming larger in swollen portion; fine ridges in blind tip; approx. 75% enclosed in penial tunica. Epiphallus enters penis through simple pore; epiphallus longer than penis; epiphallus 2 longer than epiphallus 1; long epiphallic caecum present, with medial attachment of the penial retractor muscle; flagellum long, slender, tightly coiled, with tiny internal cryptae. Remarks Pravonitor ferrugineus was introduced for a medium-sized, banded snail from the Iron Range (Stanisic et al. 2010). It was described as differing from P. kreffti by its flatter shell and the presence of a strong supraperipheral reddish-brown band. We found two morphospecies in material identified as P. ferrugineus, similar in genital morphology but differing in the size, shape and internal sculpture of the penis and in their flagellum (and presumably spermatophore) shape. In particular, there is a marked difference in the penis interior, which is sculptured with very fine longitudinal pilasters in one species and with pointed, conical pustules in the other species. Both had pale cream shells with a strong reddish-brown band, differing slightly in size and shape. However, the sequences available (a single sequence from each morphospecies) showed a genetic divergence of only 2.5%. Given the very distinct differences seen in the genitalia, we treat these morphospecies as distinct despite the low genetic differentiation. Through comparison with photographs of the holotype, we identified the smaller, higher-spired shell as P. ferrugineus. The second species is described below as P. stuarti sp. nov. Pravonitor ferrugineus is recorded only from the Iron Range, Far North Qld, where it is found in rainforest, in and under logs (Stanisic et al. 2010). Herein we expand the known distribution to include Heathlands National Park, 120 km to the north (Figure 4). It is possible that P. ferrugineus is sympatric with P. stuarti, which is recorded from High Range, only 30 km south-east of the type locality of P. ferrugineus, but at present there is insufficient data to clarify this. Another population with the same distinctive colouring as P. ferrugineus and P. stuarti is known: Helicarionidae CY 7 from Black Mountain, south of Cooktown. The only material available for examination was a single adult shell and a spirit-preserved juvenile specimen, so species identity could not be confirmed. Pravonitor ferrugineus can be distinguished from most of its congeners and by members of Geminitor and Malandena by its cream-coloured shell with a distinct reddishbrown supraperipheral band, and from P. stuarti by its smaller size and higher, conical rather than concave spire., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1780-1782, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press."]}

Geminitor laura (Stanisic, 2010) (Figures 4, 6 (a), 7(a–c), 8(a), 9) Pravonitor laura Stanisic, 2010: 300–301. Material examined Holotype. QM MO53669, Palmerville Stn, SSE on Maytown Rd, Far North Qld, Australia (16.047°S, 144.214°E), among limestone rock, 17 December 1996, L. Little. Paratypes. QM MO61204, QM MO61357, QM MO61431, QM MO61701, QM MO61763, QM MO62896, QM MO78932, AM C.425417. Other material. See Table 1. Diagnosis External morphology. Shell (Figures 6 (a), 7(a–c)) medium-sized (SW 11.6–14.6 mm), golden amber, 5.5–6.3 whorls, discoidal with a low to moderately elevated spire, whorls rounded; umbilicus narrow, sealed; protoconch with moderately coarse spiral grooves, becoming finer and more crowded on teleoconch. Body (Figure 8 (a)) pale brown with grey tail tip and black eyestalks. Mantle lobes small; shell lappets moderately short, narrow, tapering. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 9) with very short vagina; bursa copulatrix moderately long, duct broad, internally with transverse ridges, bursa elongate. Penis large, internally with two chambers, distal portion sculptured with two distinct longitudinal pilasters; proximal portion sculptured with transverse rounded plates arranged like petals around opening to epiphallus; approx. 75–100% enclosed in penial tunica. Epiphallus enters penis through a small, fragile verge; epiphallus longer than penis, epiphallus 2 similar in width to penis, gradually narrowing, epiphallus 1 narrow; boundary between penis and epiphallus 2 indistinguishable; epiphallus 1 slightly longer than epiphallus 2; long epiphallic caecum present, with basal attachment of the penial retractor muscle; flagellum moderately long, slender, without obvious internal cryptae. Remarks Geminitor laura was described from Palmerville, and ranges north to the Laura Basin and east to near Cooktown, Far North Qld, in dry vine thicket and adjacent woodland (Stanisic et al. 2010). We extend the range herein to include populations from farther south at Mt Elephant (Figure 4). Considerable genetic variation was observed within this species, as with some other northern Qld helicarionids that exhibit broad ranges (e.g. P. aquilonia). However, anatomical differences were slight at most, and given the paucity of data, we choose to retain all populations in G. laura at the present time. Geminitor villaris, G. macveae, and G. cf kullaensis are found in very close proximity to G. laura, although no sympatry is recorded. Geminitor laura can be distinguished from these species by its significantly larger size, and anatomically by the lack of a penial verge in either chamber of the penis., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1758-1762, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press."]}

Geminitor villaris (Reeve, 1854) (Figures 4, 6 (b–c), 8(b), 10) Helix villaris Reeve, 1854: pl. 195, sp. 1375; Pfeiffer 1855: 146; Cox 1864: 15. Xesta villaris: Cox 1868: 2, pl. 10. fig. 8. Helix (Thalassia) villaris: Brazier 1876: 118. Nanina (Macrochlamys) villaris: Pfeiffer 1878 –1881: 44. Nanina (Subg. Xesta Sect. Macrochlamys) villaris: Tryon 1886: pl. 105, pl. 35, fig. 41. Pravonitor villaris: Stanisic, 2010: 300–301, 326. Material examined Syntypes. NHMUK 1982243, Lizard Is., Qld, Australia, 3 specimens. Other material. See Table 1. Diagnosis External morphology. Shell (Figure 6 (b–c)) small (SW 8.1–10.7 mm), pale gold, 4.8– 5.7 whorls, subdiscoidal with a moderately elevated spire, whorls rounded; umbilicus narrow, sealed; protoconch with moderately coarse spiral grooves, becoming finer and more crowded on teleoconch. Body (Figure 8 (b)) white with grey on dorsal part of tail and tail tip, and black eyestalks (in alcohol). Mantle lobes small; shell lappets triangular, tapering. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 10) with short, swollen vagina; bursa copulatrix moderately short, duct broad, bursa rounded. Penis large, internally with two chambers, distal portion with thin outer wall and thicker inner wall, sculptured with a few longitudinal ridges, and a single large, ridged longitudinal pilaster with a caecum-like extension; proximal portion sculptured with transverse elongate plates arranged like petals around penial verge; 100% enclosed in penial tunica. Epiphallus enters penis through a small, beehive-shaped verge; epiphallus longer than penis, epiphallus 2 similar in width to penis, gradually narrowing, epiphallus 1 slightly narrower; epiphallus 2 longer than epiphallus 1; long epiphallic caecum present, with basal attachment of the penial retractor muscle; flagellum moderately short, slender, without obvious internal cryptae. Spermatophore simple, consisting of soft capsule and long, slender tail-pipe sculptured with two ridges with indistinct teeth. Remarks This species was described as Helix villaris from Lizard Island by Reeve (1854), who noted the delicate, pale-green horny shell of five whorls, with a minute umbilicus and very finely sutured margin. No measurements were given, but the most mature syntype specimen is approximately 10 mm in width. The species was attributed to Pfeiffer (1855) by Reeve and all contemporary authors, but Reeve’s publication precedes Pfeiffer’s and it is now accepted as the original description. Pfeiffer gave the type locality as ‘Cape York, Torres Strait, Lizard Point, Australia’ rather than Lizard Island, and subsequently the species was for some years regarded as coming from much farther north. It was briefly synonymised with Pravonitor kreffti from Cape York (Cox 1868), and was recorded by Brazier (1876) from Cape York and Albany Island following the Chevert Expedition. These records probably refer to the species described herein as Pravonitor septentrionalis. Iredale (1937) relegated this species to a junior synonym of Expocystis rustica, followed by Smith (1992) and Hyman and Ponder (2010). However, Stanisic et al. (2010) reinstated Helix villaris, placing it in the genus Pravonitor. Herein we have placed this species in new genus Geminitor, based on anatomical characters including the presence of a two-chambered penis, as well as genetic data. No topotype material exists other than the syntypes, which are merely empty shells. We have examined the available spirit-preserved material and found that it appears to represent four different species. One lot (QM MO23884) we tentatively assigned to G. kullaensis (see Remarks for G. kullaensis below). A second (QM MO20694), referred to above as P. villaris ‘Rifle Creek’, was smaller than the recorded measurement of P. villaris and is found significantly farther south, and is described below as the new species Geminitor macveae sp. nov. The remaining material fell into two categories: multiple lots from around Cooktown (referred to above as P. villaris ‘Cooktown’), contrasting with a single lot from 50 km farther north, from Starcke Station Gate near Mt Webb National Park (P. villaris ‘Starcke’). Both were very similar in external appearance. Their internal anatomy was also very similar, indicating a very close relationship if not conspecificity, but showed some consistent differences when several specimens from each population were compared. The Starcke specimens had a more elongate reproductive system, with a longer and slenderer penis, epiphallus and flagellum (Figure 10 (d–f)). The upper chamber of the penis was much longer, and the penial verge appeared longer and more slender as well. Both species had a single large longitudinal pilaster, folded and wrinkled, but in the Cooktown specimens this was larger and had a caecum-like extension (not attached to the penial wall). It is not clear whether these differences are sufficient to ensure reproductive isolation. Moreover, without topotype material, and with such a strong similarity between the populations, it is very difficult to determine which should bear the name P. villaris. Therefore, we elect to retain both populations in P. villaris at the present time, with a recommendation for further collecting and genetic studies. Geminitor villaris is recorded from north of the Bloomfield River to Cape Melville and off-lying islands, in dry vine thicket and adjacent woodland (Stanisic et al. 2010). Our studies can only confirm a smaller range, from Cooktown to Mt Webb National Park (Figure 4). This species is very similar anatomically to its parapatric congener G. laura but differs in its significantly smaller size and paler shell and body colouration. It also has a more distinct, swollen vagina, and differs in the internal anatomy of the penis and its shorter bursa copulatrix and flagellum., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1762-1765, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Reeve LA. 1854. Monograph of the genus Helix, part II. In: Reeve LA, editor. Conchologia Iconica: or, illustrations of the Molluscous animals. Vol. VII. London: Lovell Reeve.","Pfeiffer L. 1855. Descriptions of fifty-seven new species of Helicea, from Mr Cuming's collection. Proc Zool Soc Lond. 1854: 286 - 298. doi: 10.1111 / j. 1469 - 7998.1854. tb 07277. x","Cox JC. 1864. Catalogue of specimens of the Australian land shells in the collection of James C. Cox. Sydney: John Alex Engel. doi: 10.5962 / bhl. title. 12929.","Cox JC. 1868. A monograph of Australian land shells. Sydney: William Maddock. doi: 10.5962 / bhl. title. 1292.","Brazier J. 1876. List of land shells collected during the Chevert expedition. Proc Linn Soc NSW. 1: 117 - 133. doi: 10.5962 / bhl. part. 12392","Tryon GW jnr. 1886. Manual of conchology; structural and systematic. Second series: pulmonata. Volume 2. Zonitidae. Philadelphia: G. W. Tryon, jnr.","Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press.","Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1."]}

Elatonitor Stanisic, 2010 Malandena Iredale, 1933: 56 [nom. nud.]. Malandena Iredale, 1937: 7 [nom. nud.]. Malandena Hyman and Ponder, 2010: 67 [nom. nud.]. Malandena Stanisic, 2010: 296 [nom. nud.]. Elatonitor Stanisic, 2010: 298 (of masculine gender). Type species Elatonitor montanus Stanisic, 2010. Differential diagnosis External appearance. Shell medium-sized to large (SW 13.9–20 mm), pale amber to amber, often deep amber below periphery, glossy, 5.3–5.8 whorls, depressedly globose to trochoidal with a moderate spire, whorl profile mildly to strongly angulate. Protoconch with very fine spiral grooves, sometimes becoming obsolete; teleoconch with very fine and dense to coarse, distinct spiral grooves. Body light to dark brown. Mantle lobes small to moderately small; shell lappets long, narrow, tapering. Caudal horn moderately large to large. Genital anatomy. Ovotestis of 8–9 lobes, embedded in digestive gland. Carrefour visible; talon usually embedded in albumen gland. Spermoviduct not folded. Free oviduct long, with elongate capsular gland; free oviduct internally sculptured with longitudinal pilasters; capsular gland with no internal sculpture. Vagina short to long, sculptured internally with longitudinal pilasters or a network of wavy ridges. Bursa copulatrix inserted on the vagina, moderately short to moderately long, duct narrow, sometimes slightly broader at base, internally with primarily longitudinal sculpture; bursa elongate oval. Penis moderately small to long, tubular; internally with 2–4 longitudinal pilasters which sometimes merge or divide, internal wall sculptured with longitudinal wrinkles or ridges; penis and sometimes part of epiphallus enclosed in penial tunica. Epiphallus enters penis through a simple pore or a short verge; epiphallus longer than penis; epiphallus 2 equal to or longer than epiphallus 1; epiphallic caecum absent; flagellum long, slender, without obvious internal cryptae. Spermatophore a soft-walled capsule with hard tail-pipe; capsule very elongate, with elongate tip; tail-pipe short, sculptured with short hooks, becoming more dense towards tip. Remarks Elatonitor was introduced for a single species, E. montanus, based on the sharply angulate whorls and very fine, very dense spiral sculpture of its shell. In the current study we demonstrate that there is a close relationship between E. montanus and Malandena suturalis, sole remaining member of genus Malandena. However, Malandena, while in current usage, has never been validly described and therefore remains unavailable (Köhler and Bouchet 2020). Consequently, Elatonitor takes its place. Like all names ending in ‘nitor’, the genus name Elatonitor is of masculine gender. Elatonitor is characterised by its medium-sized to large, glossy shell (shell 15–20 mm in width), depressedly globose to trochoidal shape with an angulate periphery and usually a supraperipheral band, and by anatomical features including the presence of at least a short vagina, a long, slender penis with longitudinal pilasters and longitudinal ridges or wrinkles, and an epiphallus with no caecum and with a slender flagellum. In particular, it can be distinguished from the similar genera Pravonitor and Geminitor by its large size, angulate shell periphery, the lack of an epiphallic caecum and the single-chambered penis with primarily longitudinal sculpture. This genus is found in rainforest and vine thicket from Mt Bartle Frere to Cooktown (Stanisic et al. 2010)., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1790-1791, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press.","Iredale T. 1933. Systematic notes on Australian land shells. Rec Aust Mus. 19 (1): 37 - 59. doi: 10.3853 / j. 0067 - 1975.19.1933.690.","Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1.","Kohler F, Bouchet P. 2020. On unavailable genus-group names introduced by Tom Iredale for Australian non-marine gastropods: nomenclatural clarifications and descriptions of new genera. Molluscan Res. 40 (2): 150 - 159. doi: 10.1080 / 13235818.2020.1724603."]}

Elatonitor suturalis (Odhner, 1917) (Figures 4, 7 (g–i), 8(f), 22(b), 24) Macrochlamys suturalis Odhner, 1917: 81, figs 35b, 36–38. Macrochlamys suturalis depressa Odhner, 1917: 81. Malandena suturalis: Iredale 1933: 56; Iredale 1937 c: 7; Smith, 1992: 236; Hyman and Ponder 2010: 67–68, figs 19 K, 20 J, 21 K, 22E, 24A, 25A–C; Stanisic et al. 2010: 296– 297, 326. Synonymy follows Smith (1992). Material examined Lectotype. NHRM SMNH Type-9346. Malanda, N Qld, Australia. Paralectotypes. NHRM 950 (Malanda, N Qld, 3 specimens), NHRM 6315 (Cedar Creek, N Qld, 2 specimens), NHRM 6316 (Mt Bellenden Ker, N Qld, 1 specimen), NHRM 6317 (Herberton, N Qld, 2 specimens). Syntype of Macrochlamys suturalis depressa. NHRM 939. Cairns and Atherton, N Qld, 1 specimen. Diagnosis External morphology. Shell (Figures 7 (g–i), 22(b)) medium-sized to large (SW 15.3–20 mm), pale amber, often deep amber below periphery, coloured reddish brown at suture and around umbilicus, 5.3–5.8 whorls, depressedly globose with a moderately raised spire, whorl profile rounded above and below a mildly angulate periphery; microsculpture of very fine (almost obsolete) spirals on protoconch, and coarser, more distinct spirals on teleoconch; umbilicus very narrow. Body (Figure 8 (f)) dark brown with black eyestalks. Mantle lobes moderately small; shell lappets long, narrow, tapering. Caudal horn large. Genital anatomy. Genitalia (Figure 24) with short vagina, internally sculptured with a network of wavy ridges; bursa copulatrix moderately long, duct narrow, slightly broader at base, internally with wavy transverse ridges at base, becoming longitudinal pilasters connected by tiny transverse ridges in most of duct; bursa oval. Penis long, tubular; internally with 2–4 longitudinal pilasters which sometimes merge or divide; internal wall sculptured with longitudinal ridges; penis enclosed in penial tunica. Epiphallus enters penis through a short verge; epiphallus longer than penis; epiphallus 2 approximately equal to epiphallus 1; epiphallic caecum absent; flagellum long, slender, without obvious internal cryptae. Spermatophore a soft-walled capsule with hard tail-pipe; capsule very elongate, with elongate tip; tail-pipe short, sculptured with short hooks, becoming more dense towards tip. Remarks In the original description of Elatonitor suturalis, this species was placed in the genus Macrochlamys and syntypes were designated from four separate locations in northern Qld (Malanda, Bellenden Ker, Cedar Creek and Herberton) (Odhner 1917). The syntype material was examined in the current study, and the specimen that had been dissected and figured by Odhner (1917) is herein designated as a lectotype in accordance with Art. 74 of the International Code of Zoological Nomenclature (ICZN 1999). This restricts the type locality to Malanda. The assignment of specimens to E. suturalis in the current study was confirmed by comparison with the dissected lectotype. Odhner’s (1917) figure agrees with the lectotype, but he mistakenly identifies the bursa copulatrix as the dart sac. Elatonitor suturalis is recorded from rainforest and vine thicket from Cardwell Range to the Atherton Tablelands (Stanisic et al. 2010); this range is confirmed in the present study (Figure 4). At the type locality, Malanda, it appears that both Pravonitor aquilonia and P. monteitshi are sympatric with E. suturalis. Both species bear a strong resemblance to E. suturalis, but can be distinguished by their much finer teleoconch sculpture, in contrast to the distinct and rather coarse spiral grooves seen on both the upper and lower shell surfaces in E. suturalis., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1792-1796, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Odhner NHJ. 1917. Results of Dr. E. Mjobergs Swedish scientific expeditions to Australia 1910 - 1913.","Iredale T. 1933. Systematic notes on Australian land shells. Rec Aust Mus. 19 (1): 37 - 59. doi: 10.3853 / j. 0067 - 1975.19.1933.690.","Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1.","Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press.","ICZN. 1999. The international code of zoological nomenclature. 4 th ed. London: The International Trust for Zoological Nomenclature."]}

Hyman, Isabel T., Köhler, Frank (2022): Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae). Journal of Natural History 56 (41-44): 1727-1799, DOI: 10.1080/00222933.2022.2136017, URL: http://dx.doi.org/10.1080/00222933.2022.2136017

Geminitor wenlockensis sp. nov. (Figures 4, 6 (f), 13) Etymology Named for the Wenlock River, one of Australia’s last remaining wild rivers. Much of the range of this species lies in the Wenlock River drainage sub-basin, with specimens (including the holotype) collected near several different river crossing points. Material examined Holotype. QM MO86000, Wenlock R, c. 17.7 km N, on Peninusla Rd, Far North Qld, Australia (−12.3, 142.6), 18 September 1988, J. Stanisic, D. Potter, J. Chaseling. Paratypes. QM MO21419 (same data as holotype). Other material. See Table 1. Diagnosis External morphology. Shell (Figure 6 (f)) small (SW 8–10 mm), pale amber, 5.5–6.1 whorls, subdiscoidal with a moderately elevated spire, whorls rounded; umbilicus narrow, sealed; protoconch with moderately coarse spiral grooves, becoming finer and more crowded on teleoconch. Body white (in alcohol). Mantle lobes small; shell lappets moderately large, triangular, tapering. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 13) with short, slightly swollen vagina; bursa copulatrix moderately long, duct broad, internally with broad longitudinal pilasters basally, followed by transverse ridges and then wavy longitudinal pilasters, bursa oval. Free oviduct very long. Penis large, internally with two chambers, distal portion sculptured with two longitudinal pilasters, one large and raised, one flat; inner penial wall sculptured with longitudinal ridges distally, transverse ridges proximally; proximal portion sculptured with transverse wavy ridges arranged like petals around opening to epiphallus; distal portion longer than proximal portion; 40% enclosed in penial tunica. Epiphallus enters penis through small, beehive-shaped verge; epiphallus approx. equal in length to penis, epiphallus 2 similar in width to penis, gradually narrowing, epiphallus 1 narrower; epiphallus 1 longer than epiphallus 2; long epiphallic caecum present, with basal attachment of the penial retractor muscle; flagellum moderately short, slender, without obvious internal cryptae. Remarks This species was preliminarily identified as Helicarionidae CY 9 based on curatorial work. Geminitor wenlockensis ranges from the Portland Rd crossing of Wenlock River to Weipa in the west and Bramwell Junction in the north (Figure 4). Some specimens of G. wenlockensis were misidentified in museum collections as the helicarionids Expocystis rusticus and Pravonitor kreffti. Expocystis rusticus is similar in appearance to G. wenlockensis, but can be distinguished by its much smaller size. It is also found farther south than G. wenlockensis, ranging from Cardwell to north of Cooktown and west to Mt Carbine and Chillagoe (Stanisic et al. 2010). Pravonitor kreffti is similar in size and appearance, but can be distinguished from G. wenlockensis by its much larger size, and is also found farther north, from Cape York to the Inner and Near Western Torres Strait Islands. There is a stronger similarity between G. wenlockensis and P. septentrionalis, another Cape York species, which is more similar in size but differs in its less glossy shell surface, unsealed umbilicus and more defined columellar thickening., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1769-1771, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press."]}

Pravonitor kreffti (Cox, 1864) (Figures 4, 14 (e–f), 18) Helix kreffti Cox, 1864: 21. Helix (Thalassia) kreffti: Brazier 1876: 118. Nanina (Subg. Xesta Sect. Macrochlamys) kreffti: Tryon 1886: 219, pl. 63, figs 98–99. Nanina (Macrochlamys) kreffti: Cox 1909: 6. Nitor (Pravonitor) kreffti: Iredale 1937: 3. Pravonitor kreffti: Smith, 1992: 241; Stanisic et al. 2010: 300–301 (in part). Material examined Holotype. Status unknown, whereabouts unknown presumed lost, Cape York, Far North Qld, Australia. Neotype. QM MO86003. Green Hill, Thursday Island, Torres Strait, Far North Qld, Australia (−10.58, 142.28), 11 April 1985, J. Stanisic, D. Potter. Paraneotype. QM MO15572 (same data as neotype). Other material. See Table 1. Diagnosis External morphology. Shell (Figure 14 (e–f)) medium-sized (SW 12.5–13.7 mm), golden brown, 5.0–5.7 whorls, subdiscoidal to subglobose with a rounded whorl profile; shell microsculpture with very fine to obsolete spiral grooves; umbilicus very narrow, sealed with a mucous plug, partially covered by reflected lip. Body grey with a cream sole (in alcohol). Right and median mantle lobes moderately small, left lobe small; shell lappets moderately large, narrow, tapering, right lappet longer than left. Caudal horn moderately large. Genital anatomy. Genitalia (Figure 18) with short vagina, internally with broad longitudinal pilasters; bursa copulatrix moderately long, duct broad at base, then narrowing, internally with broad longitudinal pilasters; bursa elongate. Penis large, narrow at base and becoming broader proximally, with a large blind tip; internally with no pilasters or with one pustulose longitudinal pilaster, internal wall sculptured with fine pustules; approx. 50–75% enclosed in penial tunica; penis sometimes folded longitudinally inside tunica. Epiphallus enters penis through a simple pore; epiphallus similar in length to penis; epiphallus 2 longer than epiphallus 1; moderately short epiphallic caecum present, with medial or apical attachment of the penial retractor muscle; flagellum long, slender, without obvious internal cryptae. Remarks The correct identity of this species requires some clarification. It was first described by Cox (1864) based on a single specimen collected by MacGillivray, with no figure and with only a brief description (including the dimensions diam. maj. 0.57, min. 0.49, alt 0.31 (uncertain), 6 whorls), and a rather general type locality of simply ‘Cape York’. Shortly afterwards, Cox (1868) synonymised the species with P. villaris, and figured the specimen under this name, with dimensions (width 0.5 inches; height 0.25 inches, shell 5 whorls) matching the figure given at natural size. Here, Cox (1868) gives the locality as ‘Cape York (Cox); Rocky Isles, near Cape Flattery (MacGillivray)’; however, this second locality is probably referring to G. villaris, which was described from Lizard Island. Subsequent authors including Brazier (1876), Tryon (1886) and Cox (1909) treated kreffti and villaris as separate species. Brazier (1876) stated that he believed Cox’s type shell to come from Darnley Island in the Torres Strait, given the close similarity of the kreffti type to the Darnley Island species,and recorded villaris from Cape York and Albany Island (this latter reference was probably actually P. septentrionalis, since G. villaris is found considerably farther south). Among material from Cape York, there are two species represented: one is relatively small (shells of 5.0–5.5 whorls are 7.8–9.3 mm in width) and is referred to above as P . kreffti ‘Cape York’. The other is much larger (shells of 5.3–5.8 whorls are 12.9–15.3 mm in width) and is conspecific with a group found primarily on the Inner and Near Western Torres Strait Islands, referred to above as P . kreffti ‘Inner/ Western Islands’; only a single lot from Cape York is known (AM C.425474 from Simpson’s Bay, Utingu, coll. 1912). Cox’s description, dimensions and figure match the larger species and we accept this as P . kreffti. The holotype was recorded as lost by Smith (1992), and cannot be found in the collections of AM or NHMUK. To clarify the taxonomic status of P . kreffti, we designate QM MO86003 from Waiben Island as a neotype in accordance with Art. 75 of the International Code of Zoological Nomenclature (ICZN 1999). This specimen was dissected and sequenced and used as a basis for our taxonomic description. It matches the dimensions of the holotype, and among the available ethanol-preserved specimens, is geographically closest to the original type locality. In accordance with Art. 76 of the Code, Green Hill, Thursday (= Waiben) Island becomes the type locality of this species. The distribution of Pravonitor kreffti was previously recorded from Weipa, east to Portland Roads and north to Bamaga on Cape York, and also the Torres Strait Islands (Stanisic et al. 2010). It is now limited to the very tip of Cape York, and the Near Western and Inner Islands of the Torres Strait (Figure 4), including Waiben, Muralug, Mua and Badu Islands, and probably numerous additional islands including Ngurupai (= Horn), Maururra (= Wednesday), Gealug (= Friday) and Kiriri (= Hammond) Islands. Populations from islands in the Central and Top Western groups and the southern coast of New Guinea are provisionally referred to P. annulus, and populations from the Eastern Islands (including Mer Island) are placed in P. insularum. Pravonitor kreffti can be distinguished from P. annulus by its slightly larger size, higher spire and rounded rather than subangulate whorl profile, and by the lack of a supraperipheral band., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1782-1784, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Cox JC. 1864. Catalogue of specimens of the Australian land shells in the collection of James C. Cox. Sydney: John Alex Engel. doi: 10.5962 / bhl. title. 12929.","Brazier J. 1876. List of land shells collected during the Chevert expedition. Proc Linn Soc NSW. 1: 117 - 133. doi: 10.5962 / bhl. part. 12392","Tryon GW jnr. 1886. Manual of conchology; structural and systematic. Second series: pulmonata. Volume 2. Zonitidae. Philadelphia: G. W. Tryon, jnr.","Cox JC. 1909. A list of the subclass Pulmonata found in Australia, Part I., not including Tasmania, Lord Howe's Island (under the NSW Government), or the New Guinea Mainland and adjacent islands. Sydney: Frederick W. White.","Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408.","Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press.","Cox JC. 1868. A monograph of Australian land shells. Sydney: William Maddock. doi: 10.5962 / bhl. title. 1292.","ICZN. 1999. The international code of zoological nomenclature. 4 th ed. London: The International Trust for Zoological Nomenclature."]}

Pravonitor Iredale, 1937 Pravonitor Iredale, 1937: 3. Type species Nitor kreffti insularum Iredale, 1937 (by original designation). Differential diagnosis External appearance. Shell medium-sized to large (SW 10–21 mm), pale gold to amber, sometimes with an amber supraperipheral band, lower half of shell often slightly darker in colour, glossy, 5–6.4 whorls, subdiscoidal to subglobose, whorls subangulate to rounded. Protoconch and teleoconch with very fine spiral grooves, sometimes becoming obsolete on teleoconch. Body colour cream to brown (in alcohol). Mantle lobes moderately small; shell lappets long, triangular, wide at base, tapering to a point. Caudal horn large. Genital anatomy. Ovotestis of 4–8 lobes, embedded in digestive gland. Carrefour visible; talon usually embedded in albumen gland. Spermoviduct not folded. Free oviduct long, with elongate capsular gland; free oviduct internally sculptured with longitudinal pilasters. Vagina short, sculptured internally with longitudinal pilasters. Bursa copulatrix moderately short to moderately long, inserted on the vagina, base of duct usually swollen, then narrowing; bursa elongate. Penis short and broad to long and slender, internally pustulose, with or without longitudinal pilasters, or with longitudinal pilasters only (P. ferruginea only). Penis tunica attached by muscle fibres to middle of epiphallus; epiphallus enters penis through simple pore or small verge; epiphallus 2 longer than epiphallus 1; epiphallic caecum moderately long to long, penial retractor muscle attached to middle of caecum. Long, slender epiphallic flagellum with axial filaxment present; internal cryptae generally minor and not visible externally (except in P. annulus). Spermatophore a softwalled capsule with hard tail-pipe; capsule with elongate tip; tail-pipe smooth, or sculptured with fine, hair-like spines. Remarks Pravonitor was first introduced as a subgenus of Nitor for two species from Far North Qld, Helix kreffti Cox, 1864 and Nitor kreffti insularum Iredale, 1937 (Iredale 1937), on the basis of their rounded rather than angulate or keeled shell periphery and their ‘different’ shell texture. Pravonitor was raised to generic status by Smith (1992), who also synonymised P. kreffti and P. kreffti insularum. Hyman and Ponder (2010) demonstrated that Pravonitor could also be clearly separated from Nitor based on anatomical data. Most recently, Stanisic et al. (2010) included in this genus Helix villaris Reeve, 1854 (previously considered a synonym of Expocystis rustica Iredale 1937; Smith, 1992) and two new species, Pravonitor laura and Pravonitor ferrugineus. In the current study we have determined that P. laura and P. villaris are not closely related to the type species of this genus, P. insularum (herein removed from synonymy with P. kreffti) but instead group with two undescribed species from Far North Qld. These two species are henceforth placed in the new genus Geminitor. In addition to P. insularum, P. kreffti and P. ferrugineus, we include in Pravonitor two species previously belonging to Malandena, M. aquilonia Stanisic, 2010 and M. monteithi Stanisic, 2010. We also describe P. septentrionalis from Cape York and P. stuarti from High Range, and reinstate P. annulus, previously listed as incertae sedis (Smith et al., 2002), bringing the total species count for this genus to eight. Pravonitor can be characterised by its medium-sized to large, highly glossy shell, with very fine to obsolete spiral microsculpture, often with a peripheral band, subdiscoidal to subglobose in shape, with a rounded to subangulate whorl profile. Anatomically, defining characters include the short vagina, elongate free oviduct, long, slender bursa copulatrix with a basally broadened duct, long epiphallic caecum with medial attachment of the penial retractor muscle, slender flagellum, sometimes with distinct internal cryptae (most pronounced in P. annulus), and generally pustulose penis interior. The spermatophore consists of a soft, elongate capsule with a long tail-pipe, sometimes smooth (P. monteithi), sometimes sculptured with hook-like spines (short in P. kreffti, longer in P. annulus). In P. annulus, both the flagellum and the spermatophore tail-pipe form a spiral., Published as part of Hyman, Isabel T. & Köhler, Frank, 2022, Homoplasy in shells discombobulated the taxonomy: revision of the larger helicarionid land snails of northern Queensland, Australia (Stylommatophora: Helicarionidae), pp. 1727-1799 in Journal of Natural History 56 (41 - 44) on pages 1771-1772, DOI: 10.1080/00222933.2022.2136017, http://zenodo.org/record/7380681, {"references":["Iredale T. 1937. A basic list of the land Mollusca of Australia. Part II. Aust Zool. 9: 1 - 39.","Cox JC. 1864. Catalogue of specimens of the Australian land shells in the collection of James C. Cox. Sydney: John Alex Engel. doi: 10.5962 / bhl. title. 12929.","Smith BJ 1992. Non-marine Mollusca. In: Houston W, editor. Zoological catalogue of Australia (Vol. 8). Canberra: Australian Government Publishing Service; pp. 1 - 408.","Hyman IT, Ponder WF. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa. 2462 (1): 1 - 148. doi: 10.11646 / zootaxa. 2462.1.1.","Stanisic J, Shea M, Potter D, Griffiths O. 2010. Australian land snails. 1. A field guide to eastern Australian species. Riviere des Anguilles (Mauritius): Bioculture Press.","Reeve LA. 1854. Monograph of the genus Helix, part II. In: Reeve LA, editor. Conchologia Iconica: or, illustrations of the Molluscous animals. Vol. VII. London: Lovell Reeve.","Smith BJ, Reid S, Ponder WF (2002) Pulmonata. Australian Faunal Directory. Australian Biological Resources Study, Canberra. Available online at http: // www. environment. gov. au / biodiversity / abrs / online-resources / fauna / afd / taxa / pulmonata [viewed on 2021 Sept 29]."]}

Thai limestone karsts are known to contain a rich biodiversity of animals, especially terrestrial snails, but still require further intensive exploration to evaluate their biodiversity. To date, only a few studies on the limestone karst-inhabiting land snail genera have been published. The present work focuses on the species diversity and phylogenetic relationships of the limestone karst-restricted land snail genus Aenigmatoconcha from Thailand, based on comparative morphology and molecular evidence. The results yielded three known species (A. clivicola Tumpeesuwan & Tumpeesuwan, 2017, A. sumonthai Tumpeesuwan & Tumpeesuwan, 2018, and A. mitis (Pfeiffer, 1863) comb. nov.), plus a new species (A. eunetis Pholyotha & Panha sp. nov). The phylogenetic analyses of partial fragments of the mitochondrial cytochrome oxidase c subunit I (COI) gene confirmed the monophyly of all recognized species and congruence with the traditional morphology-based species designations. Average uncorrected p-distances of COI sequences between species were 9.7–12.0% and within species were 0.2–4.2%. This study also provides the re-description of penial sculpture, penial sheath, flagellum, penial caecum, and mantle lobe morphology that were neglected from the type species description. The present discovery of a new species increases the known diversity of Thai land snails and will support the conservation planning to protect karst biodiversity.

Salween River basin’s karst ecosystems in southeastern Myanmar remain largely unexplored and are likely to harbour a high terrestrial snail diversity that are often associated with high levels of snail endemism. Here, an outstanding group of new karst-associated terrestrial snails, Burmochlamysgen. nov., are discovered. A study of the comparative morphological and anatomical data reveals that the reproductive tract and radula of this new genus are closely related to the helicarionid genus Sophina Benson, 1859 but shell morphology (shape, size, and sculpture) and mantle extensions are distinct from the latter genus. Burmochlamysgen. nov. now consists of four known nominal species, B. cassidulacomb. nov., B. cauisacomb. nov., B. perpaulacomb. nov., and B. poongeecomb. nov., and five new species; B. albidasp. nov., B. fasciolasp. nov., B. moulmeinicasp. nov., B. versicolor sp. nov., and B. whittenisp. nov. The highlight is that the members of the new genus show site-specific endemism, being restricted to karstic habitat islands of the Salween River basin. In addition, the discovery supports that the unique and complex structure of Salween River basin’s karst ecosystems are habitats in which the terrestrial malacofauna have speciated and become endemic.

This study presents a list of land snails and slugs found on limestone hills in the District of Bau, the state of Sarawak in Malaysian Borneo. Systematic and random sampling for land snails was conducted at eight limestone outcrops, namely, Gunung Stulang, Padang Pan, Gunung Kapor, Gunung Lobang Angin, Gunung Doya, Gunung Batu, Bukit Sekunyit and Gunung Sebayat. A total of 122 land snail species was documented with photographs of each species. Of the 122 species collected, 13 are new to science, namely,Acmella bauensissp. nov.,Japonia bauensissp. nov.,Plectostoma margaretchanaesp. nov.,Microcystina arabiisp. nov.,Microcystina atonisp. nov.,Microcystina pariparisp. nov.,Microcystina liratasp. nov.,Microcystina oswaldbrakenisp. nov.,Microcystina kilatsp. nov.,Philalanka jambusanensissp. nov.,Everettia microrhytidasp. nov.,Everettia minutasp. nov., andParalaoma sarawakensissp. nov.

Aenigmatoconcha eunetis Pholyotha & Panha sp. nov. urn:lsid:zoobank.org:act: 98AD360F-93A3-4A8E-BE1A-5C991DBC358F Figs 1, 2F–H, 7E–F, 9, 10D Diagnosis Shell medium-sized and pale yellowish white. Aperture ovate-lunate in shape and vertically open. Genitalia with very long and slender penis with many tiny conical penial pilasters inside. Radular teeth arranged in wide-angle U-shaped row, teeth with oblong shape, monocuspid. Etymology The specific name ʻ eunetis ʼ is from the Greek word meaning ʻspousesʼ, honouring the authors of genus Aenigmatoconcha, who are married. Material examined Holotype THAILAND • 1 sh (width 13.9 mm, height 7.4 mm); Uthai Thani Province, Lan Sak District, Limestone outcrops at Tham Namthip Bureau of Monks; 15°26′00.3″ N, 99°35′18.7″ E; CUMZ 7931. Paratypes THAILAND • 1 sh, 7 sp; same collection data as for holotype; CUMZ 7933 • 12 sp; same collection data as for holotype; CUMZ 7932 • 2 sh; same collection data as for holotype; NHMUK. Other material THAILAND • 1 sp; Uthai Thani Province, Lan Sak District, Limestone outcrops at Hup Pa Tat; 15°22′36.5″ N, 99°37′49.5″ E; CUMZ 7934 • 6 sp; same collection data as for preceding; CUMZ 7935 • 3 sh, 27 sp; Uthai Thani Province, Nong Chang District, Limestone outcrops at Wat Khao Bang Kraek; 15°18′07.9″ N, 99°41′04.5″ E; CUMZ 7936. Description SHELL (Fig. 7E–F). Depressed to globosely depressed, medium-sized (width 12.6–14.2 mm, height 6.1– 7.1 mm), thin and translucent. Shell surface smooth, and polished, and pale yellowish white. Whorls: 5–5½, regularly increasing in size, separated by shallow suture. Spire rather elevated; varix usually present; last whorl well-rounded. Aperture very obliquely oval-lunate in shape; peristome simple. Columellar margin simple and slightly expanded near umbilicus. Umbilicus open and deep. EXTERNAL FEATURES (Fig. 2F–H). Living snails with reticulated skin and pale yellowish to slightly dark grey body. Five well-developed mantle lobes: left and right shell lobes translucent, same colour as body, covered by tiny whitish dots, right shell lobe smaller than left shell lobe. Dorsal lobes broad and crescent-shaped: right dorsal lobe larger than both anterior and posterior left dorsal lobes. Caudal fossa present; caudal horn raised and rather large, and same colour as body. GENITALIA (Fig. 9). Atrium (at) enlarged and very short. Penis (p) very long and slender with rather thick penial sheath (ps) covering from atrium to almost middle of penis. Inner sculpture of penis with small conical penial pilasters (pp), and three prominent longitudinal folds along entire length of penis chamber. Epiphallus (e1 + e2) approximately half of penis length: e1 elongate and slender, and e2 very short and bulbous. Inner sculpture of e1 nearly smooth with small, thin, and longitudinal folds, and e2 densely papillate. Epiphallic caecum (ec) straight, short, and approximately as long as e2; penial retractor muscle (prm) thin and attached at tip. Flagellum (fl) small, rather short, and approximately as long as e2. Vas deferens (vd) very long, thin, and convoluted. Vagina (v) short, enlarged, thickened and cylindrical. Gametolytic sac (gs) short and bulbous; gametolytic duct (gd) long, enlarged near vagina then becoming smaller and very slender. Free oviduct (fo) long, cylindrical, and encircled with dense tissue near vagina. RADULA (Fig. 10D). Teeth arranged in wide-angle U-shape with half row consisting of about 97–98 teeth at middle plate. Central tooth symmetrical monocuspid, elongated oblong-shaped with curved cusp, and slightly smaller than lateral tooth. Lateral and marginal teeth undifferentiated, asymmetrical monocuspid and elongated oblong-shaped with curved cusp; outermost teeth gradually becoming smaller. Distribution Aenigmatoconcha eunetis sp. nov. occurs in a few isolated limestone hills in Uthai Thani Province.This new species lives on limestone karsts, where snails tend to hide themselves in rock crevices and shelter during the daytime. Remarks Aenigmatoconcha eunetis sp. nov. clearly differs from all congeners in having 1) the longest penis, 2) penial internal sculpture consisting of longitudinal folds and small conical penial pilasters, and 3) inner sculpture of epiphallus (e2) densely papillae. For comparison, A. clivicola has oblique trapezoidshaped penial pilasters, A. mitis has oblique wrinkled penial pilasters with a longitudinal fold, and A. sumonthai has very small conical penial pilasters without longitudinal fold. In addition, for the inner sculpture of epiphallus (e2), A. clivicola has irregularly oblique trapezoid and small conical papillae. Aenigmatoconcha mitis and A. sumonthai have loose papillae arranged in oblique rows, but the former has relatively fewer rows and papillae are significantly larger in size than the latter. Moreover, radular teeth of only A. eunetis sp. nov. are arranged in wide-angle U-shaped row and oblong-shaped teeth, while radular teeth of other Aenigmatoconcha species are arranged in V-shaped rows and spatulate in shape. Discussion Each of the four Aenigmatoconcha species forms a well-defined clade in the COI phylogeny. Aenigmatoconcha can be divided into two groups based on the genital characters (Figs 1–2; Table 2). Group I (Fig. 2) has a long epiphallic caecum and short penial caecum, and contains one species, A. clivicola, which is restricted to northeasthern Thailand (Fig. 1). Group II (Fig. 2) has a short epiphallic caecum and no penial caecum; this group contains A. mitis, A. sumonthai, and A. eunetis sp. nov. This group occurs from central to southern Thailand along the Tenasserim Range (Fig. 1). The COI tree showed very low nodal support for the relationships among the two Aenigmatoconcha groups and the other two helicarionid genera. Although the phylogenetic relationships among Aenigmatoconcha, Sophina and Chalepotaxis remain unresolved, the genital characters of both groups of Aenigmatoconcha were clearly distinct from Sophina and Chalepotaxis. In the phylogenetic tree of the 28S gene, moreover, these three genera were confirmed as different genera, but the systematic position of two Aenigmatoconcha species (A. clivicola and A. sumonthai) is unresolved (Sutcharit et al. 2020a). Further research should include on more genes and more taxa of the Southeast Asian helicarionoids to better understand the phylogenetic relationships and morphological evolution of these groups. Aenigmatoconcha can be distinguished from almost all other Southeast Asian helicarionoid genera by their unique milky to pale whitish-horny and umbilicate shell. Other helicarionoids with a whitish shell include Macrochlamys psyche Vermeulen et al., 2019, Sarika lactoconcha Pholyotha & Panha, 2020, and Sarika consepta (Benson, 1860). However, the relatively medium to large-sized shells (shell width larger than 15 mm), narrowly perforate umbilicus, genital structure, and radula with triangular teeth of these three species clearly differentiate them from all Aenigmatoconcha species (Vermeulen et al. 2019; Pholyotha et al. 2020 a, 2020b). The greatest similarity in shell characters occurred in Sophina and Chalepotaxis, but their genitalia are obviously different. The distinctive characters between Aenigmatoconcha and Chalepotaxis have been reported (Tumpeesuwan & Tumpeesuwan 2017; Sutcharit et al. 2020a). However, Tumpeesuwan and Tumpeesuwan (2017) did not report the presence of a “penial sheath” and a “flagellum” in A. clivicola (the type species). In fact, they stated that the absence of a penial sheath in Aenigmatoconcha is a diagnostic difference between Aenigmatoconcha and Chalepotaxis. Yet, in the present work, the four Aenigmatoconcha species clearly exhibited a well-developed penial sheath and flagellum. Hence, the only difference between Chalepotaxis and Aenigmatoconcha is the presence of a flagellum in the latter (Páll-Gergely et al. 2016). In addition to the genitalia without a dart apparatus, the presence of well-developed anterior and posterior left dorsal lobes is a significant character of Aenigmatoconcha, while Sophina has an undivided left dorsal lobe and a dart apparatus (Blanford & Godwin-Austen 1908; Sutcharit et al. 2020a). However, information on the mantle lobe morphology of Chalepotaxis was not available for comparison. Among Southeast Asian helicarionoid genera with uniform and spatulate-shaped radula, Sophina is the only genus that a dart apparatus present in genitalia, but a flagellum absent. While genitalia of Aenigmatoconcha lack dart apparatus, but contain the flagellum, and genitalia of Chalepotaxis lack both organs. Generally, some characters of genital anatomy, such as a penial caecum, penial verge, dart apparatus, or flagellum, have been hypothesized to evolve repeatedly during the evolution of land snails, and have been noticed in many groups of terrestrial pulmonate snails (Solem 1966; Hausdorf 1998; Hyman & Ponder 2010; Hirano et al. 2014; Köhler & Criscione 2015; Köhler et al. 2020; Sutcharit et al. 2020a). However, the classification of these three genera based on the presence or absence of the dart apparatus and flagellum is consistent with the molecular phylogeny (Sutcharit et al. 2020a). From the East of Tenasserim Range to northeastern Thailand, three Aenigmatoconcha species occurred in a few limestone karsts, while only A. mitis occurs in many limestone karsts from central to western Thailand (Fig. 1). Aenigmatoconcha clivicola is confined to Loei Province, northeastern Thailand, while A. eunetis sp. nov. to Uthai Thani Province, central Thailand, and A. sumonthai to Chumphon Province, southern Thailand. The isolation explains the degree of endemism and the very high genetic divergence among sister lineages (9.7% to 12.0%) within Aenigmatoconcha. Regarding the West of Tenasserim Range, Sophina also reveals a high degree of endemism and localization with a pattern of one outcrop for one lineage in the Salween Basin, Southern Myanmar. These phenomena can be generally observed in karst-restricted animals because a very large number of ecological niches in karst ecosystems promote their evolutionary diversification and evolution of remarkably different lifestyles (Clements et al. 2006; Foon et al. 2017; Grismer et al. 2020; Sutcharit et al. 2020a). Therefore, our discovery enhances the understanding of karst biodiversity and supplements the information on terrestrial snails in Thailand available for efforts to establish well-planned and knowledge-based conservation procedures for Thai limestone protection in the future., Published as part of Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 2021, Systematic revision of the limestone karst-restricted land snail genus Aenigmatoconcha (Eupulmonata: Helicarionidae), with description of a new species, pp. 55-82 in European Journal of Taxonomy 767 on pages 72-77, DOI: 10.5852/ejt.2021.767.1487, http://zenodo.org/record/5528134, {"references":["Sutcharit C., Jeratthitikul E., Pholyotha A., Lin A. & Panha S. 2020 a. Molecular phylogeny reveals high diversity and endemism in the limestone karst-restricted land snail genus Sophina Benson, 1859 from Myanmar (Eupulmonata: Helicarionidae), with description of four new species. Journal of Zoological Systematics and Evolutionary Research 58: 957 - 981. https: // doi. org / 10.1111 / jzs. 12420","Vermeulen J. J., Luu H. T., Theary K. & Anker K. 2019. New species of land snails (Mollusca: Gastropoda: Caenogastropoda and Pulmonata) of the Mekong Delta Limestone Hills (Cambodia, Vietnam). Folia Malacologica 27: 7 - 41. https: // doi. org / 10.12657 / folmal. 027.001","Pholyotha A., Sutcharit C., Tongkerd P. & Panha S. 2020 b. Integrative taxonomic revision of the land snail genus Sarika Godwin-Austen, 1907 in Thailand, with descriptions of nine new species (Eupulmonata, Ariophantidae). ZooKeys 976: 1 - 100. https: // doi. org / 10.3897 / zookeys. 976.53859","Pholyotha A., Sutcharit C., Thach P., Chhuoy S., Ngor P. B. & Panha S. 2020 a. Land snail genus Sarika Godwin-Austen, 1907 (Eupulmonata: Ariophantidae) from Cambodia, with description of three new species. European Journal of Taxonomy 674: 1 - 21. https: // doi. org / 10.5852 / ejt. 2020.674","Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598","Pall-Gergely B., Feher Z., Otani J. U. & Asami T. 2016. An integrative taxonomic approach to infer the systematic position of Chalepotaxis Ancey, 1887 (Gastropoda: Stylommatophora: Helicarionidae). Molluscan Research 37: 113 - 119. https: // doi. org / 10.1080 / 13235818.2016.1234996","Pfeiffer L. 1863. Descriptions of thirty-six new land shells from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 30: 268 - 278. [Published in parts, dates follow Duncan (1937)].","Solem A. 1966. Some non-marine mollusks from Thailand, with notes on classification of the Helicarionidae. Spolia Zoologica Musei Hauniensis 24: 1 - 110.","Hausdorf B. 1998. Phylogeny of the Limacoidea sensu lato (Gastropoda: Stylommatophora). Journal of Molluscan Studies 64: 35 - 66. https: // doi. org / 10.1093 / mollus / 64.1.35","Hyman I. T. & Ponder W. F. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa 2462: 1 - 148. http: // doi. org / 10.11646 / zootaxa. 2462.1.1","Hirano T., Kameda Y., Kimura K. & Chiba S. 2014. Substantial incongruence among the morphology, taxonomy, and molecular phylogeny of the land snails Aegista, Landouria, Trishoplita, and Pseudobuliminus (Pulmonata: Bradybaenidae) occurring in East Asia. Molecular Phylogenetics and Evolution 70: 171 - 181. https: // doi. org / 10.1016 / j. ympev. 2013.09.020","Kohler F. & Criscione F. 2015. A molecular phylogeny of camaenid land snails from northwestern Australia unravels widespread homoplasy in morphological characters (Gastropoda, Helicoidea). Molecular Phylogenetics and Evolution 83: 44 - 55. https: // doi. org / 10.1016 / j. ympev. 2014.11.009","Kohler F., Criscione F. & Shea M. 2020. A mitochondrial phylogeny uncovers taxonomic ambiguity and complex phylogeographic patterns in the eastern Australian land snail Austrochloritis (Stylommatophora, Camaenidae). Journal of Zoological Systematics and Evolutionary Research 58: 1005 - 1020. https: // doi. org / 10.1111 / jzs. 12366","Clements R., Sodhi N. S., Schilthuizen M. & Ng P. K. 2006. Limestone karsts of Southeast Asia: Imperiled arks of biodiversity. Bioscience 56 (9): 733 - 742. https: // doi. org / fq 9 mjv","Foon J. K., Clements G. R. & Liew T. - S. 2017. Diversity and biogeography of land snails (Mollusca, Gastropoda) in the limestone hills of Perak, Peninsular Malaysia. ZooKeys 682: 1 - 94. https: // doi. org / 10.3897 / zookeys. 682.12999","Grismer L. L., Wood P. L., Le M. D., Quah E. S. H. & Grismer J. L. 2020. Evolution of habitat preference in 243 species of Bent-toed geckos (Genus Cyrtodactylus Gray, 1827) with a discussion of karst habitat conservation. Ecology and Evolution 10: 13717 - 13730. https: // doi. org / 10.1002 / ece 3.6961"]}

Aenigmatoconcha mitis (Pfeiffer, 1863) comb. nov. Figs 1, 2B, 4C–F, 6, 10B Helix mitis Pfeiffer, 1863 [1862]: 268. Type locality: Lao Mountains, Camboja [Lao Mountains, Cambodia]. Helix mitis – Pfeiffer 1868: 141. — Tryon 1886: 171. Ariophanta (Kaliella) mitis – Fischer 1891: 21. Hyalinia mitis – Fischer & Dautzenberg 1904: 396. Macrochlamys (?) mitis – Inkhavilay et al. 2019: 78, 79, fig. 37b. Material examined Syntypes CAMBODIA • 2 sh; “Lao Mountains, Camboja ” [Lao Mountains, Cambodia]; NHMUK ex. Cuming collection. Other material THAILAND • 37 sh, 16 sp; Nakhon Sawan Province, Mueang District, Limestone outcrops at Wat Tham Bo Ya; 15º43′47.3″ N, 99º56′44.7″ E; CUMZ 7708 • 30 sh, 15 sp; same collection data as for preceding; CUMZ 7885 • 8 sh, 21 sp; Nakhon Sawan Province, Krok Phra District, Limestone outcrops at Wat Khao Tham Phra; 15°33′30.2″ N, 99°57′28.1″ E; CUMZ 7849 • 3 sp; Kanchanaburi Province, Sai Yok District, Limestone outcrops at Wat Thep Thepa Satthatham; 14°03′56.6″ N, 99°11′45.7″ E; CUMZ 7913 • 21 sh, 24 sp; Kanchanaburi Province, Mueang District, Limestone outcrops at Wat Tham Charoentham; 13°55′59.1″ N, 99°27′59.9″ E; CUMZ 7687 • 3 sp; Kanchanaburi Province, Tha Muang District, Limestone outcrops at Wat Tham Faet; 13°57′52.5″ N, 99°34′56.1″ E; CUMZ 7914 • 30 sh, 1 sp; Kanchanaburi Province, Dan Makham Tia District, Limestone outcrops at Wat Tham Khao Cha Ang; 13°48′08.4″ N, 99°26′33.2″ E; CUMZ 7247 • 4 sh, 3 sp; Phetchaburi Province, Khao Yoi District, Limestone outcrops at Wat Khiri Wong; 13°20′03.2″ N, 99°45′19.0″ E; CUMZ 7917 • 1 sh, 15 sp; Phetchaburi Province, Khao Yoi District, Limestone outcrops at Wat Puang Malai; 13°18′46.0″ N, 99°47′02.2″ E; CUMZ 7921 • 9 sh; same collection data as for preceding; CUMZ 7938 • 5 sh, 14 sp; Phetchaburi Province, Tha Yang District, Limestone outcrops at Wat Khao Krachiu; 12°57′41.3″ N, 99°54′49.3″ E; CUMZ 7918 • 3 sh, 16 sp; Phetchaburi Province, Cha-am District, Limestone outcrops at Khao Nang Panthurat; 12°50′20.5″ N, 99°57′11.6″ E; CUMZ 7920 • 2 sh, 2 sp; Phetchaburi Province, Cha-am District, Limestone outcrops at Tham Chaeng Bureau of Monks; 12°49′44.3″ N, 99°56′27.1″ E; CUMZ 7919 • 4 sh, 7 sp; Prachuap Khiri Khan Province, Mueang District, Limestone outcrops at Khao Lom Muak; 11°47′03.0″ N, 99°48′56.3″ E; CUMZ 7915 • 3 sh, 8 sp; Prachuap Khiri Khan Province, Mueang District, Limestone outcrops at Khao Ta Mong Lai; 11 o 50′07.3″ N, 99 o 49′50.1″ E; CUMZ 7916. Description SHELL (Fig. 4C–F). Shell depressed to globosely depressed, medium-sized (width 12.3–18.1 mm, height 6.9–9.9 mm), rather thin, and translucent. Shell surface smooth and glossy. Shell colour whitish to very pale horny-white. Whorls: 5–5½, regularly increasing in size; varix present; suture rather wide and shallow. Spire rather elevated. Last whorl broad and well-rounded. Aperture obliquely oval-lunate in shape; peristome simple. Columellar margin simple, slightly expanded near umbilicus. Umbilicus open and deep. EXTERNAL FEATURES (Fig. 2B). Living snails with reticulated skin and pale yellowish to dark grey body. Five well-developed mantle lobes; left and right shell lobes thin, pale yellowish, spread with small whitish dots, and left shell lobe larger than right shell lobe. Three dorsal lobes broad and crescentshaped; right dorsal lobe larger than anterior and posterior left dorsal lobes. Caudal fossa present; caudal horn raised, rather large, and whitish to pale fleshy-grey in colour. GENITALIA (Fig. 6). Atrium (at) enlarged and very short. Penis (p) long, cylindrical, and with slightly thick penial sheath (ps) encircling about half of penis length. Inner sculpture of penis with very small and oblique wrinkled penial pilasters (pp), and one large longitudinal fold running the length of the entire penis chamber. Epiphallus (e1 + e2) approximately as long as penis: e1 slender and narrower than penis, and e2 shorter and bulbous shape. Inner sculpture of e1 with very small thin longitudinal folds to nearly smooth surface with one thickened longitudinal fold, and inner sculpture of e2 with large papillae arranged in oblique rows. Epiphallic caecum (ec) very short; penial retractor muscle (prm) thin and attached at tip. Flagellum (fl) small and rather short. Vas deferens (vd) very long and thin. Vagina (v) very short and enlarged. Gametolytic duct (gd) long, slender, and enlarged near vagina; gametolytic sac (gs) very large and oblong shape. Free oviduct (fo) cylindrical, long, and encircled with thick tissue near vagina. RADULA (Fig. 10B). Teeth arranged in anteriorly V-shaped rows with half row consisting of about 76– 79 teeth at middle plate. Central teeth symmetrical monocuspid, and spatulate-shaped with curved cusp. Lateral and marginal teeth undifferentiated, asymmetrical monocuspid, spatulate-shaped with curved cusp, and outermost teeth gradually reduced in size. Distribution The distribution of Aenigmatoconcha mitis is wider than all other recognised species. This species can be found in limestone areas ranging from central (Nakhon Sawan Province) to southern (Prachuap Khiri Khan Province) Thailand (Fig. 1). Remarks This species was originally described by L. Pfeiffer (1863) based on specimens in the collection of H. Cuming obtained from Henry Mouhot. The collection locality was brief: “Lao Mountains, Camboja ”. However, Mouhot’s recorded localities were generally imprecise and referred to a wide geographical area, for example “ Siam ”, “Lao Mountains, Camboja ” and “ Camboja ”. This has made it difficult to infer more precise type localities of several land snail species described from Mouhot’s specimens. No additional specimen records or literature references are available for this species until now. The most recent works on land snails from Laos and southern Cambodia confirmed the existence of A. mitis (Inkhavilay et al. 2019; Sutcharit et al. 2020c). Based on the recorded itinerary, H. Mouhot had travelled to “Pechaburi” [Petchaburi Province] in 1861, and clearly stated that he had visited caves and several hills during his four-month stay (Mouhot 1864: 57; Ashburton 1864: map). We have surveyed several limestone hills in western and peninsular Thailand and encountered numbers of empty shells and living specimens that well-matched with the type specimens of “ Helix mitis Pfeiffer, 1863 ” (Fig. 4C). Therefore, peninsular Thailand (Petchaburi Province; Fig. 1) might be the area where H. Mouhot collected this species. Aenigmatoconcha mitis exhibits a rather wide range of shell shape variation from depressed (Fig. 4F) to somewhat globose (Fig. 4C, D). However, the genitalia of these shell morphs are identical and the COI phylogeny also supports that all shell morphs are grouped together within the A. mitis clade (Fig. 2)., Published as part of Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 2021, Systematic revision of the limestone karst-restricted land snail genus Aenigmatoconcha (Eupulmonata: Helicarionidae), with description of a new species, pp. 55-82 in European Journal of Taxonomy 767 on pages 66-70, DOI: 10.5852/ejt.2021.767.1487, http://zenodo.org/record/5528134, {"references":["Pfeiffer L. 1863. Descriptions of thirty-six new land shells from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 30: 268 - 278. [Published in parts, dates follow Duncan (1937)].","Pfeiffer L. 1868. Monographia heliceorum viventium supplementum tertium: sistens descriptiones systematicas et criticas omnium hujus familiae generum et specierum hodie cognitarum, volume quintum. F. A. Brockhaus, Lipsiae.","Tryon Jr. G. W. 1886. Manual of Conchology; Structural and Systematic. With illustrations of the species. Second Series: Pulmonata. Volume II. Zonitidae. Academy of Natural Sciences, Philadelphia.","Fischer P. 1891. Catalogue et distribution geographique des Mollusques terrestres, fluviatiles & marins dune partie de lIndo-Chine (Siam, Laos, Cambodge, Cochinchine, Annam, Tonkin). Imprimerie Dejussieu Pere et Fils, Autun. https: // doi. org / 10.5962 / bhl. title. 14809","Fischer H. & Dautzenberg P. 1904. Catalogue des Mollusques terrestres et fluviatiles de lIndo-Chine orientale cites jusqu' a ce jour. Mission Pavie, Etudes diverses 3: 390 - 450. https: // gallica. bnf. fr / ark: / 12148 / bpt 6 k 8814596 [accessed 22 July 2021]","Inkhavilay K., Sutcharit C., Bantaowong U., Chanabun R., Siriwut W., Srisonchai R., Pholyotha A., Jirapatrasilp P. & Panha S. 2019. Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda). ZooKeys 834: 1 - 166. http: // doi. org / 10.3897 / zookeys. 834.28800","Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598","Sutcharit C., Thach P., Chhuoy S., Ngor P. B., Jeratthitikul E., Siriwut W., Srisonchai R., Ng T. H., Pholyotha A., Jirapatrasilp P. & Panha S. 2020 c. Annotated checklist of the land snail fauna from southern Cambodia (Mollusca, Gastropoda). ZooKeys 948: 1 - 46. https: // doi. org / 10.3897 / zookeys. 948.51671","Mouhot H. 1864. Travels in the Central Parts of Indo-China (Siam), Cambodia, and Laos, During the Years 1858, 1859, and 1860. Murray, London.","Ashburton J. 1864. Paper read at the Royal Geographical Society 10 th March, 1862. In: Mouhot H. (ed.) Travels in the Central Parts of Indo-China (Siam), Cambodia, and Laos, During the Years 1858, 1859, and 1860: 296 - 300. Murray, London."]}

Genus Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017 Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017: 182–184. Aenigmatoconcha – Tumpeesuwan & Tumpeesuwan 2018: 171. Type species Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, by original designation. Description Shell dextral with 5–5½ convex whorls, strongly depressed to globosely depressed, medium-sized, thin to slightly solid, translucent, and pale milky to whitish-horny in colour. Shell surface smooth, glossy, and varix usually present. Body whorl well-rounded to slightly-shouldered. Suture shallow. Aperture slightly to very crescentic in shape with simple lip. Umbilicus open and deep. Animal with reticulated skinand whitish, yellowish, pale fleshy grey to dark brown body with tiny whitish dots irregularly scattered over entire body. Mantle lobes well-developed (two shell lobes and three dorsal lobes; see Figs 3, 5A). Shell lobes can cover most, if not all of the shell and are retracted when disturbed. Left and right shell lobes very thin, translucent, ovate to triangular in shape; right shell lobe (rsl) smaller than left shell lobe (lsl). Dorsal lobes enlarged, crescent-shaped, covering body, and smaller than shell lobes. Anterior left dorsal lobe (ant-ldl) and posterior left dorsal lobe (post-ldl) smaller than right dorsal lobe (rdl). Sole tripartite, lateral foot margin, caudal fossa, and caudal horn present. Genitalia with moderately long to very long penis, thick penial sheath, short to long epiphallus, small flagellum, and short vagina. Gametolytic organ with short gametolytic duct and bulbous gametolytic sac. Oviduct with large lobules; prostate gland running alongside oviduct. Radular teeth arranged in anteriorly V-shaped or wide-angle U-shaped rows; central tooth symmetrical monocuspid and spatulate or oblong in shape; lateral and marginal teeth undifferentiated, asymmetrical monocuspid and spatulate or oblong in shape, and outermost teeth gradually reduced in size. Distribution Species of Aenigmatoconcha exhibit allopatric distributions and are restricted to limestone karsts in Thailand (Fig. 1)., Published as part of Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 2021, Systematic revision of the limestone karst-restricted land snail genus Aenigmatoconcha (Eupulmonata: Helicarionidae), with description of a new species, pp. 55-82 in European Journal of Taxonomy 767 on pages 62-64, DOI: 10.5852/ejt.2021.767.1487, http://zenodo.org/record/5528134, {"references":["Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598"]}

Aenigmatoconcha sumonthai Tumpeesuwan & Tumpeesuwan, 2018 Figs 1, 2C–E, 7A–D, 8, 10C Aenigmatoconcha sumonthai Tumpeesuwan & Tumpeesuwan, 2018: 171–173, figs 2–6. Type locality: Tham Chang Phueak limestone range, Mueang District, Chumphon Province, southern Thailand. Material examined THAILAND • 45 sh, 20 sp; Chumphon Province, Mueang District, Limestone outcrops at Tham Chang Phuek Bureau of Monks; 10°26′50.0″ N, 99°02′07.1″ E; CUMZ 7922 • 40 sh; same collection data as for preceding; CUMZ 7923 • 18 sh, 10 sp; same collection data as for preceding; CUMZ 7937 • 45 sh, 6 sp; Chumphon Province, Mueang District, Limestone outcrops at Wat Tham Sanook; 10°28′51.3″ N, 99°04′28.3″ E; CUMZ 7924 • 10 sh, 7 sp; same collection data as for preceding; CUMZ 7925 • 22 sh, 11 sp; Chumphon Province, Sawi District, Limestone outcrops at Tham Nam Lod Thepnimit Bureau of Monks; 10°22′39.5″ N, 99°00′39.5″ E; CUMZ 7927 • 4 sh, 8 sp; Chumphon Province, Sawi District, Limestone outcrops at Wat Nam Cha; 10°17′57.0″ N, 99°01′58.5″ E; CUMZ 7926. Description SHELL (Fig. 7A–B). Shell strongly depressed to depressed, medium-sized (width 14.4–16.6 mm, height 7.0– 8.2 mm), thin, translucent, whitish colour, well-rounded to slightly shouldered body whorl, elevated spire, impressed suture, obvious varix, and open umbilicus. EXTERNAL FEATURES (Fig. 2C–E). Animal with five well-developed mantle lobes. Left and right shell lobes pale yellowish to fleshy-grey colour, usually with black margin, and with or without small to large black spots or blotches. Three dorsal lobes crescent-shaped and smaller than the other two shell lobes. Black stripes behind long tentacles. GENITALIA (Fig. 8). Atrium (at) enlarged and very short. Penis (p) long, cylindrical with thick penial sheath (ps) extending to half of penis length. Inner sculpture of penis with small conical penial pilasters (pp). Epiphallus (e1 + e2) as long as penis: e1 slender, and e2 bulbous. Inner sculpture of e1 with small thin longitudinal folds, while e2 with small papillae arranged in oblique rows. Epiphallic caecum (ec) short with thin penial retractor muscle (prm) attached at tip. Flagellum (fl) small and short. RADULA (Fig. 10C). Teeth arranged in anteriorly V-shaped rows with half row consisting of about 63– 65 teeth at the middle plate. All teeth monocuspid and spatulate-shaped with curved cusp. Distribution Aenigmatoconcha sumonthai has a narrow distribution, with populations living on a few limestone hills in Chumphon Province (Fig. 1). We extended our survey, especially among limestone sites about 200 km southwards down to southern peninsular Thailand, but we could not find this species elsewhere. Remarks The lack of a penial sheath and flagellum in the male reproductive organs of A. sumonthai was originally reported to be similar to A. clivicola (Tumpeesuwan & Tumpeesuwan, 2018: 174, fig. 6). In this study, based on topotypic specimens, however, A. sumonthai was found to have a large and thickened penial sheath and small flagellum encircled with loose tissue (Fig. 8). Aenigmatoconcha sumonthai shows variation in the black blotches on both shell lobes ranging from absent (Fig. 2E) to the lobes almost entirely covered (Fig. 2C). The DNA sequence analysis suggested that these variations formed a clade of A. sumonthai (Fig. 2). In addition, this species develops a calcareous epiphragm with a small perforation to limit body-water evaporation but allowing respiratory gas exchange during dormancy (Fig. 7C, D).

Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017 Figs 1, 2A, 4A–B, 5, 10A Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017: 184–187, figs 2–5. Type locality: Phu Pha Lom Limestone Hill in Mueang District, Loei Province, northeastern Thailand. Aenigmatoconcha clivicola – Tumpeesuwan & Tumpeesuwan 2018: 171. Material examined THAILAND • 39 sh, 22 sp; Loei Province, Mueang District, Phu Pha Lom Limestone Hill; 17°33′16.6″ N, 101°52′05.4″ E; CUMZ 7928 • 6 sh, 12 sp; same collection data as for preceding; CUMZ 7929 • 48 sh; Loei Province, Na Duang District, Limestone outcrops at Wat Tham Pha Ya; 17°34′40.1″ N, 101°53′35.1″ E; CUMZ 7930. Description SHELL (Fig. 4A–B). Shell strongly depressed to depressed, medium-sized (shell width 17.2–20.1 mm, shell height 8.6–10.0 mm), rather thin to slightly solid and translucent, milky to pale whitish-horny colour, well-rounded body whorl, little elevated spire, impressed suture, obvious varix, and open umbilicus. EXTERNAL FEATURES (Figs 2A, 5A). Animal with five well-developed mantle lobes. Left and right shell lobes thin, pale yellowish to fleshy-grey colour and spread with small whitish dots. Three dorsal lobes crescent-shaped and smaller than shell lobes. GENITALIA (Fig. 5B–D). Atrium (at) enlarged and very short. Penis (p) rather long cylindrical, penial sheath (ps) very thick and covering entire penis, and penial caecum (pc) rather small protruding. Inner wall of penis covered with trapezoid-shaped penial pilasters (pp) arranged in oblique rows. Epiphallus (e1 + e2) as long as penis: e1 long and slender, and e2 bulbous and about half e1 length. Inner sculpture of e1 with very small and thin longitudinal folds to nearly smooth surface. Inner sculpture of e2 trapezoid to conical pilasters. Epiphallic caecum (ec) long with thick penial retractor muscle (prm) attached at tip. Flagellum (fl) small and short. RADULA (Fig. 10A). Teeth arranged in anteriorly V-shaped rows with half row consisting of about 70– 76 teeth at middle plate. Central, lateral, and marginal teeth monocuspid and spatulate-shaped with curved cusp. Distribution This species is currently known only from 2 localities: the type locality (Phu Pha Lom) and limestone hills in Loei Province (Fig. 1). Remarks The genitalia were originally described but without examination of the internal sculpture of penis. In this study, we examined 20 adult topotypic specimens to provide descriptions of the penial sheath, penial caecum, and flagellum that were not included in the original description. The penial sheath is very large and covers the entire penis (Fig. 5B). The short penial caecum, an extension of the penis, is located near the penis and epiphallus junction (Fig. 5B). This character is visible when penial sheath is removed. Its internal sculpture is rather smooth, unlike the penial sculpture, with its trapezoid-shaped pilasters (Fig. 5C). The Australian helicarionids, Nitor whitneyae Stanisic, 2010 has a penial caecum while other Nitor taxa do not have a penial caecum (Hyman & Köhler 2018). In Southeast Asian ariophantids, the presence or absence of a penial caecum is a discriminating character among species in genera such as Macrochlamys Gray, 1847 and Taphrenalla Pholyotha & Panha, 2020, and is supported by molecular studies (Pholyotha et al. 2018, 2021). The flagellum of A. clivicola is an extension of the epiphallus and is located near the insertion point of the vas deferens. It is somewhat small and short, and bound to the vas deferens by thin connective tissue. This feature is important for spermatophore formation before copulation (Tompa 1984; Baur 2010). However, during this study no spermatophores were observed in A. clivicola., Published as part of Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 2021, Systematic revision of the limestone karst-restricted land snail genus Aenigmatoconcha (Eupulmonata: Helicarionidae), with description of a new species, pp. 55-82 in European Journal of Taxonomy 767 on pages 64-66, DOI: 10.5852/ejt.2021.767.1487, http://zenodo.org/record/5528134, {"references":["Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598","Pfeiffer L. 1863. Descriptions of thirty-six new land shells from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 30: 268 - 278. [Published in parts, dates follow Duncan (1937)].","Pholyotha A., Sutcharit C., Tongkerd P. & Panha S. 2020 b. Integrative taxonomic revision of the land snail genus Sarika Godwin-Austen, 1907 in Thailand, with descriptions of nine new species (Eupulmonata, Ariophantidae). ZooKeys 976: 1 - 100. https: // doi. org / 10.3897 / zookeys. 976.53859","Pholyotha A., Sutcharit C. & Panha S. 2018. The land snail genus Macrochlamys Gray, 1847 from Thailand, with descriptions of five new species (Pulmonata: Ariophantidae). Raffles Bulletin of Zoology 66: 763 - 781.","Pholyotha A., Sutcharit C., Tongkerd P., Jeratthitikul E. & Panha S. 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. https: // doi. org / 10.1163 / 18759866 - BJA 10013","Tompa A. S. 1984. Land snails (Stylommatophora). In: Tompa A. S. & Verdonk N. H. (eds) The Mollusca, Vol. 7: Reproduction: 47 - 140. Academic Press, London.","Baur B. 2010. Stylommatophoran Gastropods. In: Leonard J. L. & Cordoba-Aguilar A. (eds) The Evolution of Primary Sexual Characters in Animals: 197 - 217. Oxford University Press, USA."]}

The helicarionid fauna of southeastern to mid-eastern Queensland is dominated by a group of semislugs with moderately reduced shells belonging to genera Fastosarion, Eungarion, Stanisicarion, Dimidarion, Macularion and Hymanarion. We comprehensively revise their systematic classification using comparative morpho-anatomy and mitochondrial phylogenetics, and demonstrate that these genera combined form a well-differentiated and monophyletic radiation. In our mitochondrial phylogeny, this radiation is divided into three main clades that are statistically well supported. One clade is also well defined in terms of diagnostic morpho-anatomical characters, but we could not identify diagnostic characters for the other two clades due to considerable levels of morpho-anatomical variation. We propose accepting only two genera, Fastosarion (with junior synonyms Eungarion, Dimidarion, and Hymanarion) and Stanisicarion (with junior synonym Macularion). Both genera represent mutually monophleytic sister taxa that can consistently be distinguished by the presence or absence of a penial verge that is fused to the penial wall and by egg shape. We also synonymise Fastosarion ameyi with F. aquavitae, F. schelli with F. helenkingae, Dimidarion peterbrocki and D. slatyeri with F. alyssa, Stanisicarion virens with S. freycineti. Revised species descriptions are presented for Fastosarion alyssa, F. aquavitae, F. brazieri, F. comerfordae, F. griseolus, F. hannianus, F. helenkingae, F. mcdonaldi, F. minerva, F. paluma, F. papillosus, F. pustulosus, F. superbus, Stanisicarion aquila and S. freycineti. Nine new species, Fastosarion deensis, F. ephelis, F. insularis, F. katatonos, F. longimentula, F. rowani, F. sarina, F. tuljun and Stanisicarion wolvi are described, bringing the total number of accepted species to 24.

The land area of Laos is composed of a large variety of undisturbed habitats, such as high mountainous areas, huge limestone karsts and the lower Mekong Basin. Therefore, Laos is expected to have a high species diversity, especially for the land snails. However, with respect to research on malacology, Laos is probably the least well-researched area for land snail diversity in Indochina (including Laos) over the past few centuries. The handful of species lists have never been systematically revised from the colonial period to the present, so these classifications are outdated. Herein we present the first comprehensive annotated checklist with an up-to-date systematic framework of the land snail fauna in Laos based on both field investigations and literature surveys. This annotated checklist is collectively composed of 231 nominal species (62 ‘prosobranch’ and 169 heterobranches), of which 221 nominal species are illustrated. The type specimens of 143 species from several museum collections and/or 144 species of newly collected specimens are illustrated. There are 58 species recorded as new to the malacofauna of the country, and two new replacement names are proposed asHemiplectalanxangnicaInkhavilay and Panha,nomen novum(Ariophantidae) andChloritiskhammouanensisInkhavilay and Panha,nomen novum(Camaenidae). Four recently described species of the genusAmphidromusfrom Laos, “thakhekensis”, “richgoldbergi”, “attapeuensis” and “phuonglinhae” are synonymized with previously described species. In addition, thirteen nominal species are listed as uncertain records that may or may not occur in Laos. This annotated checklist may inspire malacologists to carry on systematic research in this region.

Anatomical study of five specimens of an endemic species of the Mauritius Island, Erepta odontina showed that within one population disappearing of additional organs of the reproductive tract (epiphallic caecum, penial caecum, penial appendix, flagellum and retractor of penis) occurs in four different combinations. It is significant that the shells of all five specimens were practically identical. It is assumed that the described phenomenon has a mutagenic nature and is regarded as a rare (if not unique) case of reduction (up to complete disappearance) of additional organs within one population.

Varadia Bhosale & Raheem gen. nov. urn:lsid:zoobank.org:act: BB363A8F-6796-4874-BBF3-6CE1A207A3C3 Type species Varadia amboliensis Bhosale, Thackeray, Muley & Raheem gen. et sp. nov. (here designated). Diagnosis Only known large semi-slug species from the Western Ghats having broad shell lobes (covering nearly all of the dorsal shell surface when fully extended) and a depressed, discoid shell with a large body whorl. Shell thin, glossy golden brown, with 4���4�� rapidly increasing whorls and barely raised spire. Extensive, largely free penial sheath enclosing substantial part of male genitalia. When sheath is in situ and penis in relaxed state, male genitalia constitute a single elongate, irregularly-shaped mass, with all of penis and epiphallic caecum, most of epiphallus and part of flagellum enclosed by sheath; only vas deferens and most of flagellum is visible outside intact sheath. Penial sheath divisible into thick proximal part and thin distal part, with transition between these two parts occurring between distal penis and epiphallic caecum. With penial sheath removed, following evident: region extending from penis to epiphallic caecum long and cylindrical, with distinctive S-shaped bend in penis and associated band of muscle running along length of penis; proximal �� of epiphallus held in long, conspicuous loop; and penial retractor muscle with two branches, one inserting subterminally on epiphallic caecum and one inserting on apex of loop of epiphallus. Inner wall of penis divisible into three regions: proximal penis (one major and several minor longitudinal pilasters), mid-penis (several minor longitudinal pilasters) and distal penis (a few minor pilasters with associated regular transverse ridges in interspaces). Inner wall of epiphallic caecum shows one long major longitudinal pilaster, a large mass of reticulate ridges proximally and several minor longitudinal pilasters distally. Gametolytic gland elongated and long; gametolytic sac 3���3.5 times as long as gametolytic duct. Amatorial organ absent. Spermatophore consists of elongated, soft capsule and long tail-pipe, with U-shaped bend at capsule���tail-pipe junction and funnel-like opening at tip of tail-pipe; surface smooth apart from four ribs running obliquely along length of tail-pipe and short spines near end of tail-pipe. Etymology The new genus is named in honour of the herpetologist Dr Varad Giri, who has made a major contribution to the modern study and conservation of the Indian herpetofauna; masculine., Published as part of Bhosale, Amrut R., Saadi, Ahmed J., Wade, Christopher M., Thackeray, Tejas U., Tamboli, Asif S., Kadam, Suhas K., Muley, Dipak V. & Raheem, Dinarzarde C., 2021, Varadia, a new helicarionoidean semi-slug genus from India's Western Ghats (Stylommatophora: Helicarionoidea), pp. 50-79 in European Journal of Taxonomy 757 (1) on page 58, DOI: 10.5852/ejt.2021.757.1413, http://zenodo.org/record/5047645

Varadia amboliensis Bhosale, Thackeray, Muley & Raheem gen. et sp. nov. urn:lsid:zoobank.org:act: 5C93F719-2DEF-4A9A-8973-06B7FE6E6528 Figs 3���12 Diagnosis As genus-level diagnosis. Etymology Named after the type locality, Amboli, in the Sindhudurg District of southern Maharashtra, India. In recent years, Amboli has emerged as a hotspot for the discovery of new species (particularly reptiles and amphibians) in the northern Western Ghats. Type material Holotype INDIA ��� Maharashtra State, Sindhudurg District, Amboli, Hiranyakeshi temple; 15��57���17.8��� N, 74��01���39.1��� E; 839 m a.s.l.; 2019; A. Bhosale leg.; BNHS GAS 113. Paratypes INDIA ��� 21 specimens (17 whole preserved specimens and 4 shells); same locality data as for holotype; 2019; A. Bhosale leg.; BNHS GAS 114���127, ZSI Moll/1820���1826 ��� 3 preserved specimens; same locality data as for holotype; 2020; A. Bhosale leg.; BNHS GAS 136���138. Other material examined INDIA ��� Maharashtra State ��� 1 specimen (sampled for DNA analysis); Sindhudurg District, Amboli Forest Park; 15��57���37.4��� N, 73��59���58.1��� E; 724 m a.s.l.; 2017; A. Bhosale leg.; BNHS GAS 129 ��� 9 preserved specimens; Sindhudurg District, near Amboli waterfall; 15��56���26.9��� N, 73��59���41.2��� E; 645 m a.s.l.; 2020; A. Bhosale leg.; BNHS GAS 130���135, BNHS GAS 139���141 ��� 1 shell; Kolhapur District, Kodali; 15��46���42.4��� N, 74��10���40.0��� E; 620 m a.s.l.; 2019; A. Bhosale leg.; BNHS GAS 128. Description SHELL.Adult shell thin, depressed, glossy and appearing non-umbilicate, with ca 4���4.5 rapidly increasing whorls and colour ranging from golden-brown to reddish yellow (Fig. 3). Shell measurements (n = 35): width 20.2���26.3 mm; height 10.2���15.0 mm. Spire only slightly raised with flat apex and suture only slightly impressed. Body whorl disproportionately large, rounded at periphery, gently convex beneath. Aperture large, crescent-shaped, with width greater than height (Fig. 3A, E). Apertural margin simple, thin and delicate; in lateral view angled forward, with upper apertural margin noticeably anterior to the lower margin. When shell is viewed from below, basal margin curved (not straight) and expanded columellar margin reflected over, covering umbilical region. Shell surface smooth and glossy to naked eye, with irregular, faint collabral striae; under SEM, seen to be finely and closely sculptured with well-defined spiral lines on protoconch (first 1.5���2 whorls) and indistinct and irregular oblique lines on teleoconch (Fig. 4). BODY AND MANTLE. Total adult body length, excluding extended tentacles, ranges from 4.8 to 6.9 cm (n = 5). Living snail glossy grey or greyish white with irregular dark mottling; head and tail dark grey or blackish with tentacles tending to be paler at their tips (Fig. 5���6). Surface of mantle densely and conspicuously covered by small, irregular tubercles that appear lighter on top. Sole tripartite with welldefined sole furrows dividing it into three distinct longitudinal tracts; central tract paler than lateral ones. Tail with large slit-like caudal pit (sensu Hausdorf 1998: 51); caudal horn prominent when extended (Fig. 6A) but when retracted gives tail truncated and blunt appearance (Fig. 6B). Mantle consists of two broad shell lobes (right and left) and two dorsal lobes (right and left) (Fig. 7). Shell lobes may cover nearly all of dorsal surface of shell (Fig. 5), but individuals have also been observed with shell lobes largely retracted and much of shell exposed (Fig. 6). Left dorsal lobe extends as far as base of tentacles when snail is resting (i.e., body not fully extended) and tentacles are retracted. REPRODUCTIVE SYSTEM. Male genitalia consist of proximally penis and distally epiphallic caecum (= epiphallic retractor caecum of Hyman & Ponder 2010: 139) and epiphallus. These three regions are held together by a penial sheath (= penial tunica of Hausdorf 1998), which is largely independent of wall of penis, epiphallic caecum and epiphallus (Figs 8A, 9A���C). Penial sheath encloses all of penis, all of epiphallic caecum, much of epiphallus and part of flagellum; it holds proximal three quarters of epiphallus in a loop against epiphallic caecum, with remaining part of epiphallus (i.e., part closest to vas deferens and flagellum) lying outside sheath, along with a substantial part of flagellum (Fig. 9A). Proximal part of penial sheath is thick and covers penis; distal half of this sheath is thin and covers epiphallus and epiphallic caecum. Thick penial sheath attached to proximal end of penis, close to genital atrium. Thin penial sheath attached to distal end of epiphallic caecum and is open where penial retractor muscle inserts on epiphallic caecum (Fig. 9A���C); an extension of the thin penial sheath also encloses a sizeable section of flagellum (this section located about halfway along length of flagellum). Epiphallus passes through and is attached to penial sheath in region where thick penial sheath transitions into thin penial sheath. With penial sheath dissected open, penis seen to have noticeable S-shaped bend midway; this bend is associated with a band of muscle that extends for some distance along penis, on either side of bend (Fig. 9D���E). Distally, penis branches into wider-lumened epiphallic caecum and narrower-lumened epiphallus. Epiphallus passes into much narrower-lumened vas deferens; junction between these two regions marked by long, bluntly pointed flagellum, which is similar in length to epiphallus. Penial retractor muscle, which originates on inner lung wall, inserts in two places (Fig. 9E): subterminally on epiphallic caecum, and on apex of loop of epiphallus (i.e., about three quarters of distance from vas deferens to penis). Junction between two branches of penial retractor muscle located near most distal part of epiphallic caecum. Irregular small holes/pores visible on inner surface of thin part of penial sheath (i.e., with sheath cut open and pinned out). On the basis of the morphology of its inner wall, penis divisible into three morphologically distinct regions, proximal penis, mid-penis and distal penis, with S-shaped bend of penis including all of mid- and distal penial regions (Fig. 10). Proximal penis shows one major and several minor longitudinal pilasters; close study at low magnification (4 ��) of holotype and one paratype (BNHS GAS 114) showed that pilasters are interspersed by fine, obliquely longitudinal ridges that are close and irregular. Mid-penis ornamented by several thin longitudinal pilasters. Distal penis also with thin longitudinal pilasters, but here they are fewer in number and are contiguous with uniform, widely spaced transverse ridges that extend outwards on either side of each pilaster. Opening of epiphallus into most proximal part of epiphallic caecum clearly visible (Fig. 10). Inner wall of epiphallic caecum (Fig. 10) has one major longitudinal pilaster (surface marked by irregular, fine longitudinal and/or transverse ridges; not shown in Fig. 10) running along its length; a large, reticulate mass of ridges proximally; and several short longitudinal pilasters distally. The short pilasters tend to be crenulated proximally and are smoother distally. Lumen of vas deferens widens with increasing distance from epiphallus, with part of vas deferens nearest to epiphallus being noticeably narrower-lumened than remaining two thirds (Fig. 8A). Right eye retractor muscle passes between male and female genitalia. Amatorial organ absent. Genital atrium cylindrical, well defined but short, with junction between male genitalia and vagina located at a short distance from genital orifice. Vagina cylindrical and shorter in length than genital atrium (Fig. 8A). Proximal part of oviduct, near junction with gametolytic gland, consists of pale yellowish, indistinctly-defined region, which is most likely the capsular gland (see Dasen 1933); inner wall of this gland irregularly marked by papillate ridges and papillae (Fig. 8). Gametolytic gland (Fig. 8A) comprises narrow gametolytic duct and long, voluminous sac that is ca 3���3.5 times length of duct; duct noticeably constricted at its junction with sac and has 1���3 longitudinal ridges on its inner wall. One or two spermatophores (i.e., only one wholly intact; the rest damaged/ partially digested) present in gametolytic gland of each of six specimens (Fig. 11A). Intact spermatophore consists of elongated, soft capsule with long tail-pipe. Sharply-angled, U-shaped bend at junction of capsule and tail-pipe; apex of bend noticeably hooked towards tail-pipe. Capsule wider-lumened than tail-pipe and twisted spirally (Fig. 11B). Tail-pipe flexible, internally hollow and externally sculptured obliquely along its length with four fine ribs. Tail-pipe in vicinity of tip hollow centrally and this passes into funnel-like opening (perforation) (Fig. 11D); surface of spermatophore near tip of tail-pipe has short, hair-like spines that point towards capsule (Fig. 11C). RADULA AND JAW. Central tooth tricuspid, with large mesocone, which is shorter than tooth base, and smaller, more basal ectocones (Fig. 12A���B). Inner laterals 17���21, uniformly tricuspid (Fig. 12A���B); mesocone large, equal in size to those of central tooth and shorter than tooth base, endocone barely defined and ectocone prominent but more basal than other cusps. Outer 2 lateral teeth grade into marginal teeth. Marginal teeth 45���53, uniformly bicuspid (endocone absent), with shorter, narrower and more basal ectocone (Fig. 12C���D). Formulae for the 8 specimens examined are as follows (the plus sign indicates that the outermost marginal teeth could not be counted): Holotype BNHS GAS 113 (+50.20.1.18.2.50+) Paratype BNHS GAS 114 (+49.21.1.19.2.49+) Paratype BNHS GAS 115 (53.19.1.17.2.53) Paratype BNHS GAS 116 (+45.21.1.19.2.45+) Paratype BNHS GAS 117 (+50.19.1.17.2.50+) Paratype BNHS GAS 118 (+48.23.1.21.2.48+) Paratype BNHS GAS 119 (+48.19.1.17.2.48+) Paratype BNHS GAS 120 (52.22.1.20.2.52) Jaw oxygnath (smooth), having a concave cutting edge with well-defined or barely evident median projection (Fig. 12E). Distribution and ecology Varadia amboliensis gen. et sp. nov. is endemic to the northern and central Western Ghats of India and is currently known from only 5 localities. These are: Hiranyakeshi temple, Amboli, Sindhudurg District, Maharashtra State (15��57���17.8��� N, 74��01���39.1��� E; 839 m a.s.l.); Amboli Forest Park, Sindhudurg District, Maharashtra State (15��57���37.4��� N, 73��59���58.1��� E; 724 m a.s.l.); near Amboli waterfall, Sindhudurg District, Maharashtra State (15��56���26.9��� N, 73��59���41.2��� E; 645 m a.s.l.); Kodali, Kolhapur District, Maharashtra State (15��46���42.4���N, 74��10���40.0���E; 620 m a.s.l.); Yana Forest, Uttara Kannada District, Karnataka State (14��35���16.4���N 74��34���00.3���E; 272 m a.s.l.) (A. Bhosale, 2018, personal observation). The species occurs at elevations ranging from 272 to 839 m. Although it has been observed among human habitation on forest edges (Fig. 13), V. amboliensis gen. et sp. nov. appears to be primarily a species of tropical semi-evergreen and evergreen forest (sensu vegetation classification of Pascal 1991). The range of this species, as currently known, is restricted and disjunct. While 4 of the 5 known localities are in the extreme south of Maharashtra State (northern Western Ghats), the only other known locality, Yana Forest in northern Karnataka (central Western Ghats), is ca 160 km to the south. Further surveys are required to establish if this species occurs in the intervening area. Varadia amboliensis gen. et sp. nov. is primarily a ground-living snail. It can be encountered at night in leaf litter or on rocks and the bases of trees; in rainy weather it can be seen on the exterior walls of buildings close to the forest edge (e.g., it was observed at the entrance of Amboli Forest Park in September 2017). The species can be seen throughout the monsoon (June to October) and as late as the end of November. A few individuals have been seen in late February (late winter) on the banks of fast-flowing streams at Amboli. This species appears to be omnivorous. It has been observed feeding on decaying plant matter (leaf litter, discarded banana peel) and on the remains of at least two different invertebrate taxa (a cricket and an earthworm) (A. Bhosale, personal observation) (Supp. file 1). Data on its predators are scarce, but a scorpion of the genus Heterometrus Ehrenberg, 1828 (Scorpionidae) was observed feeding on an individual of this species (Supp. file 1). DNA analysis The three phylogenetic analyses (NJ, ML and Bayesian) yielded broadly similar results (Fig. 14), with disagreements occurring only for internal branches lacking strong support in any of the analyses. Most deeper relationships within the Limacoidea were not strongly supported, with the optimal ML and NJ trees having fewer strongly supported branches than the Bayesian tree. All three trees included a sister- group relationship between Varadia gen. nov. and the always maximally supported Macrochlamys clade (Macrochlamys indica Benson, 1883 + M. pedina (Benson, 1865)), but this was not strongly supported in any of the analyses (Bayesian: PP = 0.62; ML: BS = 60%; NJ: BS = 44%). All analyses also provided maximal support for the clade composed of five of the six helicarionid taxa, Fastosarion brazieri (Cox, 1873), Eurychlamys platychlamys (Blanford, 1880), Harmogenanina argentea (Reeve, 1852), Plegma caelatura (F��russac, 1821) and Satiella sp. (clade B in Fig. 14). Within clade B, support for the sistergroup relationship between Fastosarion Iredale, 1933 and Eurychlamys was consistently strong (ML: BS = 90%; NJ: BS = 96%) or maximal (Bayesian). Two of the three analyses (ML: BS = 71%; Bayesian: PP = 0.98) provided strong support for clade A, comprising clade B, Rhysotina hepatzion (Gould, 1848), the Macrochlamys clade, Varadia gen. nov. and Mariaella dussumieri Gray, 1855. All analyses provided maximal support for the monophyly of the Trochomorphidae M��llendorff, 1890, of the Dyakiidae Gude & B.B. Woodward, 1921 and of the Euconulidae H.B. Baker, 1928 (i.e., clade comprising Euconulus fulvus (M��ller, 1774) and Louisia barclayi (Benson, 1850)). The monophyly of the Vitrinidae Fitzinger, 1833 and of the Oxychilidae Hesse, 1927 (1879) was also consistently strongly supported. Within the ingroup, nearly all the other branches were either strongly supported only in the Bayesian tree (e.g., clade comprising the five species of Ariophantinae sensu stricto) or were not strongly supported in any of the analyses. Examination of all bipartition frequencies for the ML bootstrap trees (n = 1008) showed that the best supported bipartition that is not compatible with Varadia gen. nov. forming a clade with the two species of Macrochlamys is one in which Macrochlamys forms a clade with the five helicarionids, Fastosarion brazieri, Eurychlamys platychlamys, Harmogenanina argentea, Plegma caelatura and Satiella sp. (BS = 12%). Similarly, for the NJ bootstrap trees (n = 1000), the best supported bipartition that is incompatible with the clade of Varadia gen. nov. + Macrochlamys is the clade uniting all the helicarionids and Rhysotina hepatzion with Macrochlamys (BS = 14%)., Published as part of Bhosale, Amrut R., Saadi, Ahmed J., Wade, Christopher M., Thackeray, Tejas U., Tamboli, Asif S., Kadam, Suhas K., Muley, Dipak V. & Raheem, Dinarzarde C., 2021, Varadia, a new helicarionoidean semi-slug genus from India's Western Ghats (Stylommatophora: Helicarionoidea), pp. 50-79 in European Journal of Taxonomy 757 (1) on pages 58-72, DOI: 10.5852/ejt.2021.757.1413, http://zenodo.org/record/5047645, {"references":["Hausdorf B. 1998. Phylogeny of the Limacoidea sensu lato (Gastropoda: Stylommatophora). Journal of Molluscan Studies 64: 35 - 66. https: // doi. org / 10.1093 / mollus / 64.1.35","Hyman I. T. & Ponder W. F. 2010. A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family. Zootaxa 2462: 1 - 148. https: // doi. org / 10.11646 / zootaxa. 2462.1.1","Dasen D. D. 1933. Structure and function of the reproductive system in Ariophanta ligulata. Proceedings of the Zoological Society of London 103: 97 - 118. https: // doi. org / 10.1111 / j. 1096 - 3642.1933. tb 01579. x","Bhosale A. R. 2018. DNA Barcoding and Phylogenetic Study of Freshwater and Land Molluscs from Kolhapur District. PhD thesis, Shivaji University, Kolhapur, India.","Pascal J. - P. 1991. Floristic composition and distribution of evergreen forests in the Western Ghats, India. Palaeobotanist 39: 110 - 126."]}

We here describe a new Indian helicarionoidean genus, Varadia Bhosale & Raheem gen. nov., containing the single species Varadia amboliensis Bhosale, Thackeray, Muley & Raheem gen. et sp. nov. This new semi-slug is endemic to the northern and central Western Ghats and is primarily a forest-living species. We describe and figure the shell, reproductive system, radula, spermatophore and external morphology of this new species, and detail its known distribution. We explore its relationships to other helicarionoideans using phylogenetic analyses of DNA sequence data for part of the ribosomal RNA gene cluster and discuss the morphology of the new genus in relation to other, primarily South Indian, helicarionoidean taxa. Based on characters of the reproductive system, particularly the male genitalia and the gametolytic sac, we provisionally place Varadia gen. nov. in the Macrochlamydinae (Ariophantidae). This is consistent with the results of our molecular phylogenetic analyses. The combination of large size, broad, densely tuberculated shell lobes and a shell with ca 4 whorls and a disproportionately large body whorl makes V. amboliensis gen. et sp. nov. unique among the helicarionoidean taxa of the Western Ghats. The new semi-slug is also highly distinctive in the morphology of its male genitalia.

Three previously unreported species of tropical land snails were found in the greenhouses of the Gothenburg (Göteborg) Botanical Garden and the Public Science Center Universeum in Gothenburg. For Tornatellides cf. boeningi (Schmacker & Boettger, 1891) and Ovachlamys fulgens (Gude, 1900) this is the first observed occurrence in a European greenhouse, while Discostrobilops hubbardi (Brown, 1861) was first reported very recently in the Vienna Botanical garden. Tornatellides and Discostrobilops seem to be spread with orchid culture and trade. Identification of the Tornatellides species proved extremely difficult and a genetic sequence-based approach completely failed due to the unavailability of reference data. This was unexpected considering the importance of these introduced species in horticultural trade. A broader assessment of available sequence data for genetic identification based on COI or 16S for other snail species reported from horticultural facilities showed that such reference data in GenBank are still scarce and only for a limited number of species this approach would support identification.

The Japanese invasive jumping snail Ovachlamys fulgens is a pest of ornamental plants and an intermediate host of a nematode that causes eosinophilic meningitis. We expand its distribution to eight municipalities from Rio de Janeiro State, and one locality from the Parana State, and generated for the first time partial sequences of the cytochrome c oxidase subunit I (COI) gene for Brazilian populations. External morphology, reproductive system, shell, radula, and jaw were also analyzed and described. Twenty-one lots were collected from Rio de Janeiro, Niteroi, Mage, Miguel Pereira, Petropolis, Teresopolis, Nova Friburgo, Bom Jardim and Paraty, in Rio de Janeiro State, and from Foz do Iguacu, Parana State. External morphology, shell and reproductive system were typical of O. fulgens, with some peculiarities found in the shell and radula. A single haplotype was found, which was 100% similar to sequences of COI available in GenBank for specimens from Japan and Argentina. The species seems to be adapted to many habitats and be rapidly expanding its distribution in Southeastern and Southern Brazil, and other South America countries. We highlight the importance of monitoring O. fulgens, considering its potential to compete with native mollusks, attack several plants, and be a transmitter of diseases.

Rosales R, Lillo E, Alcazar SM, Colita L, Caballero J, Malaki AB. 2020. Species composition, relative abundance, and distribution of land snail species in Mt. Lantoy Key Biodiversity Area, Cebu, Philippines. Biodiversitas 21: 5438-5447. This study aimed to establish an inventory of land snails in Mt. Lantoy, Cebu Island, Philippines. A two-person team conducted sampling on eight permanent plots with an elevation range of 182 m-691 m. The study has listed a total of 872 individuals represented by 25 species belonging to 15 genera and nine families. Seven families belong to the air-breathing pulmonate group, Order Stylommatophora: Bradybaenidae, Camaenidae, Trochomorphidae, Achatinidae, Ariophantidae, Helicarionidae, Helicinidae. Two families were under the prosobranch group, Order Caenogastopoda: Cyclophoridae, and Pupinidae. The Cyclophoridae was the richest family with five species, followed by Bradybaenidae, Camaenidae, Helicinidae, having four species. The Trochomorphidae and Helicarionidae have three species. The Pupinidae has two species, while the Achatinidae and Ariophantidae has one species each. There were three (3) endemic and ten (10) new island record species collected and identified with four (4) species that were classified as rare. Rysotta otaheitana Ferussac 1821 was the most abundant, representing 25.22% of the total number of individuals. The Leptoma nitidum ancilis Bartsch 1918 has the widest geographical distribution with a constancy rate of 100%. Widespread species appear to be more abundant, while other species were geographically restricted in few study areas.