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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]."]}

INNESOCONCHA PRINCEPS IREDALE, 1944 (FIGS 5J, 6H, I, 11, 22D–F, 24; TABLE 1) Innesoconcha princeps Iredale, 1944: 326, pl. 20, fig. 5; Smith, 1992: 235; Stanisic et al., 2010: 344–345; Hyman & Köhler, 2020: 102–103. Type material: Syntypes, AM C.38958 (Fig. 5J), Mount Gower (31°35.4′S, 159°4.35′E), 1912, R. Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5J) small (SW 8.8–9.1 mm, SH 4.5–5.5 mm), 4.9–6.0 whorls, glossy, light golden-brown with a darker brown peripheral stripe, appearing darker in live specimens, depressed trochoidal with a low spire. Protoconch and teleoconch sculptured with fine, incised spiral grooves. Whorls flattened above and rounded below an angulate periphery; sutures weakly impressed. Animal (Fig. 6H, I) pale to dark grey in life, faded to cream in alcohol. Right and left shell lappets narrow, finger-shaped. Caudal horn moderately large; caudal foss a vertical slit in tail. ReproductiƲe system (Fig. 24): Oviduct with 22 eggs, seven embryos. Penis long, coiled in tunica; epiphallus enters penis through a simple pore; penis internally flat, two longitudinal penis pilasters present. Penial tunica enclosing coiled penis and epiphallus; penis retractor muscle attached to penis and adjacent epiphallus. Epiphallus shorter than penis, internally with longitudinal pilasters. Spermatophore small, softwalled capsule, closed at both ends, one end rounded, one pointed. Radula (Fig. 22D–F): Central and lateral tooth mesocones approximately equal in length to tooth base. Marginal teeth multicuspidate. Radular formula (70.7.1.7.70) × 103., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 68, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 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: Bioculture Press.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

FAMILY MICROCYSTIDAE Diagnosis (based on studies by Baker, 1938; Hyman & Ponder, 2010) Shell with continuous growth and a simple, sharp peristome, rarely reduced; umbilicus closed or narrowly open; shell sculpture consisting of at most microscopic spiral grooves (most taxa) or prominent radial ribs (Fanulena Iredale, 1945 and Pittoconcha Preston, 1913). Animal with caudal apparatus formed from curled-up sole [Muratov’s (1999) helicarionid type]. Kidney unilobed; mantle with no visible minor blood vessels. Genital system ovoviviparous; oviduct non-glandular; bursa copulatrix usually very short, inserted on penis; stimulator absent; muscle fibres connecting proximal end of penial tunica to epiphallus absent; epiphallic retractor caecum and flagellum absent in most taxa. Spermatophore simple, small, peanut-shaped, soft-walled; attached to inner wall of penis. Radula with tricuspidate central tooth; lateral and marginal tooth fields distinguishable; lateral teeth usually tricuspidate (endocone absent in some taxa); marginal teeth usually multicuspidate. Penial nerve from right pedal ganglion., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 48, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Baker HB. 1938. Zonitid snails from Pacific Islands Part 1. 1. Southern genera of Microcystinae. Bernice P. Bishop Museum Bulletin 158: 3 - 102.","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 - 148.","Muratov IV. 1999. Analysis of the phylogenetic relationships and their systematic implications in the Limacoinei (= Zonitinia) infraorder (Gastropoda, Pulmonata, Geophila). Ruthenica 9: 5 - 26."]}

Annacharis miranda Iredale, 1944: 324, pl. 20, fig. 2. Melloconcha miranda Smith, 1992: 238; Hyman & Köhler, 2020: 110. Type material: Holotype, AM C.39175 (Fig. 5H), Mount Gower summit (31°35.38′S, 159°4.39′E), under stones in watercourse, 9 November 1913, W. R. B. Oliver. Material examined: See Table 1. Description Shell (Fig. 5H) small (SW 6.7 mm, SH 3.7 mm), approximately six whorls, very glossy, pale gold, globosely turbinate with a low spire. Protoconch and teleoconch sculptured with faint microscopic grooves. Whorl profile rounded above and below a rounded periphery; sutures deeply channelled on later whorls. Columella thickened., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 65, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 1 - 408.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA GRATA (IREDALE, 1944) (FIGS 5G, 6F, 13, 16G–I, 20; TABLE 1) Melloconcha grata Iredale, 1944: 323, pl. 20, fig, 13; Smith, 1992: 238; Hyman & Köhler, 2020: 109. Type material: Lectotype (herein designated), AM C.63475 (Fig. 5G), Mount Gower (31°35′23″S, 159°4′20″E), 1912, R. Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5G) small (SW 5.6– 6.6 mm, SH 3.6–4.5 mm), 4.9–5.5 whorls, not glossy, amber, depressed trochoidal. Protoconch and teleoconch with fine, incised spiral grooves. Whorl profile rounded above and below a weakly angulate periphery. Animal (Fig. 6F) grey, head and eyestalks dark grey, sole pale (faded to white in old spirit-preserved specimens). Right shell lappet long, narrow, finger-shaped; left shell lappet absent; caudal horn small. ReproductiƲe system (Fig. 20): Oviduct with one to two eggs, one to two embryos. Penis moderately long, coiled twice in tunica; epiphallus enters penis through a simple pore; penial wall with irregular longitudinal ridges, including ridges radiating out from basal region of penis; one major longitudinal penis pilaster present; two short minor pilasters apically, one short minor pilaster medially. Penial tunica enclosing coiled penis and epiphallus; penis retractor muscle attached to epiphallus. Epiphallus approximately equal to penis length, internally with longitudinal pilasters. Radula (Fig. 16G–I): Lateral teeth with mesocones slightly longer than tooth base. Marginal teeth multicuspidate. Radular formula (62.6.1.6.62) × 105 (AM C.91365)., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 64, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 1 - 408.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA DELECTA (IREDALE, 1944) (FIGS 5D, 6C, 14, 15, 16A–C) Melloconcha delecta Iredale, 1944: 323, pl. 20, fig. 8; Smith, 1992: 237; Stanisic et al., 2010: 334–335; Hyman & Köhler, 2020: 106. Type material: Holotype, AM C.63478 (Fig. 5D), North Bay (31°31.14′S, 159°2.7′E), 1912, R.Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5D) small (SW 5.3– 5.9 mm, SH 3.0– 4.3 mm), 4.6–5.8 whorls, glossy, light yellow-brown, domed in shape with a raised spire. Microsculpture of fine, incised spiral grooves. Whorl profile rounded above and below a rounded periphery; sutures slightly impressed. Animal (Fig. 6C) grey to black in life, faded to white in alcohol. Right shell lappet long, narrow, finger-shaped, left shell lappet absent; caudal horn small. Pallial caƲity: Mantle with pigmentation of white spots. ReproductiƲe system (Fig. 14): Oviduct with one to two eggs, one to two embryos. Penis short; penis not fully coiled but twisted only once at epiphallus end; epiphallus enters penis through a simple pore; penis internally flat, three to four longitudinal penis pilasters present, penial diverticulum absent. Penial tunica enclosing coiled penis and epiphallus; penis retractor muscle attached to junction of penis and epiphallus and to parts of epiphallus. Epiphallus approximately equal in length to penis, internally with longitudinal pilasters. Spermatophore a short, soft-walled, smooth, peanut-shaped tube, closed at both ends. Radula (Fig. 16A–C): Marginal teeth with ectocones approximately same length and breadth as mesocone, multicuspidate. Radular formula (54.7.1.7.54) × ~120, (46.7.1.7.46) × ~120, (45.7.1.7.45) × ~120 (all AM C.391383); (~70.7.1.7.~70) × ~100 (AM C.391761)., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on pages 59-60, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 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: Bioculture Press.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA MIRANDA (IREDALE, 1944) (FIGS 5H, 21; TABLE 1), Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 65, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334."]}

INNESOCONCHA ROSACEA (IREDALE, 1944) (FIGS 5K–O, 6J, 22G–I, 25, 26; TABLE 1) Tribocystis rosacea Iredale, 1944: 323, pl. 20, fig. 7. Melloconcha rosacea Smith, 1992: 238; Stanisic et al., 2010: 346–347; Hyman & Köhler, 2020: 112. Tribocystis alma Iredale, 1944: 324. Melloconcha alma Smith, 1992: 237. Type material: Holotype, AM C.38940 (Fig. 5K), near The Pines (31°32.08′S, 159°4.14′E), 1912, R.Bell. Holotype of Tribocystis alma, AM C. 63477 (Fig. 5L), North Ridge, between North Bay & Hunter Bay (31°31.06′S, 159°2.97′E), 1905, R.Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5K–O) small (SW 3.7–4.1 mm, SH 1.7–2.2 mm), 4.8–5.2 whorls, highly glossy, slippery, orange-brown to reddish brown, discoidal with a flat or slightly raised spire. Protoconch and teleoconch sculptured with fine microspiral grooves. Whorl profile rounded above and below a rounded periphery, sutures flat. Animal (Fig. 6J) grey, with darker grey eyestalks and tail tip. Right shell lappet long, narrow, finger-shaped, left shell lappet absent. Pallial caƲity: Pigmentation absent; mantle gland present. ReproductiƲe system (Fig. 25): Oviduct with six eggs, one to three embryos. Penis short, not coiled; epiphallus enters penis through a simple pore or small verge (present in one specimen); penis internally flat, one to two longitudinal penis pilasters present; apical diverticulum on penis present. Penial tunica enclosing penis (not coiled) and epiphallus. Epiphallus shorter than penis, internally without sculpture. Radula (Fig. 22G–I): Marginal teeth with ectocones slightly shorter and narrower than mesocone, multicuspidate. Radular formula (18.4.1.4.18) × 69., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on pages 69-70, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 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: Bioculture Press.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA ABERRANS (IREDALE, 1944) (FIGS 5C, 6B, 10G–I, 12, 13; TABLE 1) Innesoconcha aberrans Iredale, 1944: 326, pl. 20, fig. 15; Smith, 1992: 234; Hyman & Köhler, 2020: 100. Type material: Holotype, AM C.63475 (Fig. 5C), Mount Gower (31°35.4′S, 159°4.35′E), 1912, R.Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5C) small (SW 5.2– 7.8 mm, SH 4.3–4.9 mm), 4.5–5.5 whorls, not glossy, golden brown (appearing darker in live specimens), trochoidal with an elevated spire. Protoconch with fine, incised spiral grooves, teleoconch smooth. Whorl profile flattened above and below an angulate periphery. Animal (Fig. 6B) black (faded to white in old spirit-preserved specimens). Right shell lappet long, narrow, finger-shaped; left shell lappet absent. Caudal horn moderately long. ReproductiƲe system (Fig. 12): Oviduct with no more than one egg, two embryos. Penis moderately long, coiled once or twice in tunica; epiphallus enters penis through a simple pore; penial wall internally with v-shaped ridges; two major longitudinal penis pilasters present; three or four short minor pilasters apically; one moderately short minor longitudinal pilaster basally, reaching the middle of the penis. Penial tunica enclosing coiled penis and epiphallus; penis retractor muscle attached to epiphallus. Epiphallus approximately equal to penis length, internally with longitudinal pilasters. Radula (Fig. 10G–I): Lateral teeth with broad, rounded mesocones slightly longer than tooth base, and relatively small ectocones. Marginal teeth primarily bicuspidate, with reduced extra cusps becoming more prominent towards outer edge. Radular formula (60.8.1.8.60) × 124 (AM C.391773)., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 57, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 1 - 408.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

Tribocystis flaƲescens Iredale, 1944: 10, pl. 20, fig. 10. Melloconcha flaƲescens Smith, 1992: 237; Stanisic et al., 2010: 344–345; Hyman & Köhler, 2020: 107. Type material: Syntypes, AM C.40600 (Fig. 5F), North Ridge between North and Hunter Bays (31°31.06′S, 159°2.97′E), 1912, R. Bell. Two specimens. Material examined: See Table 1. Description External morphology: Shell (Fig. 5F) small (SW 5.0– 6.5 mm, SH 2.7–3.7 mm), glossy, 4.8–5.4 whorls, golden brown, discoidal with a low spire. Protoconch with fine spiral grooves; teleoconch with fine to obsolete spiral grooves. Whorl profile slightly flattened above and rounded below a rounded to slightly angulate periphery. Animal (Fig. 6D, E) cream with dark grey neck, head and eye tentacles; end of tail often with reddish, orange or yellow coloration. Right shell lappet long, narrow, finger-shaped; left shell lappet absent; caudal horn small. Pallial caƲity: Mantle with pigmentation of white spots. ReproductiƲe system (Fig. 18): Oviduct with two eggs, two to five embryos. Penis moderately short, coiled once or twice in tunica; epiphallus enters penis through simple pore; penis internally with l o n g i t u d i n a l a n d v - s h a p e d r i d g e s; t w o m a j o r longitudinal penis pilasters present; four to five minor pilasters present apically; penial diverticulum absent. Penis retractor muscle attached to junction of penis and epiphallus. Epiphallus approximately equal to penis in length, internally with longitudinal pilasters. Spermatophore a short, soft-walled, smooth, peanut-shaped tube, closed at both ends. Radula (Fig. 16D–F): Marginal teeth with ectocones approximately same length and breadth as mesocone, multicuspidate. Radular formula (56.6.1.6.56) × ~90 (AM C.399051)., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on pages 61-62, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 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: Bioculture Press.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA CATLETTI (BRAZIER, 1872) (FIGS 5A, B, 6A, 7–9, 10A–F, 11; TABLE 1) Helix (Microcystis) catletti Brazier, 1872: 617. Nanina (Microcystis) catletti: Pfeiffer, 1878 (in 1878–1881): 35. Microcystis catletti var. major Hedley, 1891: 137. Innesoconcha catletti subconica Iredale, 1944: 326 [nom. nov. for Microcystis catletti var. major Hedley, 1891]. Innesoconcha catletti Iredale, 1944: 326; Smith, 1992: 234–235; Hyman & Köhler, 2020: 101. Type material: Syntypes, AM C.101187 (Fig. 5A), AM C.63671, AM C.101064; Lord Howe Island, May 1869, G. Masters. Innesoconcha catletti major syntypes [AM C.101188 (Fig. 5B), AM C.101054], Old Settlement (31°31.16′S, 159°3.24′E), August 1887, Australian Museum party. Material examined: See Table 1. Description External morphology: Shell (Fig. 5A, B) medium-sized (SW 8.1–11.8 mm, SH 4.0– 6.8 mm), 5.0–6.1 whorls, glossy, pale golden brown, depressed trochoidal, spire and apex slightly raised. Protoconch sculptured with incised spiral grooves; teleoconch smooth. Whorl profile rounded above and below a slightly angulate periphery. Animal (Fig. 6A) cream. Right and left shell lappets narrow, finger-shaped. Caudal horn moderately small. ReproductiƲe system (Figs 7–9): Oviduct with nought to seven eggs, nought to fourteen embryos. Penis long, coiled multiple times in tunica; epiphallus enters penis through a simple pore; penis internally flat, two longitudinal penis pilasters apically and one to three pilasters present basally. Penial tunica enclosing coiled penis and epiphallus; penis retractor muscle attached to epiphallus. Epiphallus much shorter than penis, internally with longitudinal pilasters. Spermatophore short, soft-walled, smooth, peanut-shaped tube, closed at both ends. Radula (Fig. 10A–F): Central and lateral tooth mesocones approximately equal in length to tooth base. Marginal tooth ectocones approximately the same length and breadth as mesocone; multicuspidate. Radular formula (62.8.1.8.62) × 150 (AM C.399376), (69.9.1.9.69) × ~115 (AM C.389125)., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on pages 50-53, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Brazier J. 1872. Descriptions of six new species of land shells from Australia and Lord Howe's Island. Proceedings of the Zoological Society of London 1872: 617 - 619.","Pfeiffer L. 1878 - 1881. Nomenclator heliceorum ViVentium qui continentur nomina omnium hujus familiae et generum et specierum hodie cognitarum disposita ex affinitate naturali. [Posthumous work, ed. S. Clessin.] Kassel: Theodor Fischer.","Hedley C. 1891. The land and freshwater shells of Lord Howe Island. Records of the Australian Museum 1: 134 - 144.","Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 1 - 408.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA SEGNA IREDALE, 1944 (FIGS 5M, 11, 27; TABLE 1) Innesoconcha segna Iredale, 1944: 326; Hyman & Köhler, 2020: 103. Type material: Holotype, AM C.114789 (Fig. 5M), Boat Harbour (31°33.67′S, 159°5.85′E), 1912, R.Bell. Material examined: See Table 1. Description External morphology: Shell (Fig. 5M) medium-sized (SW 10.7–12.0 mm, SH 5.9–7.2 mm), 5.7–6.1 whorls, glossy, pale yellow-gold, depressed trochoidal with a low spire. Whorls rounded above and below a rounded periphery; sutures slightly impressed. Columella thickened to form a small tooth. Body colour (in alcohol) light orangebrown. Right shell lappet moderately long, narrow, fingershaped; left shell lappet short, narrow, finger-shaped. ReproductiƲe system (Fig. 27): Oviduct with four eggs, one embryo. Penis moderately long, coiled approximately twice in penial tunica; epiphallus enters penis through simple pore; penis internally with small transverse ridges, two longitudinal penis pilasters present. Penial tunica enclosing coiled penis and epiphallus. Epiphallus shorter than penis. Spermatophore short, soft-walled, smooth, peanutshaped tube, closed at both ends., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 71, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Hyman IT, Kohler F. 2020. A field guide to the land snails of Lord Hoaee Island. Sydney: Australian Museum Scientific Publishing."]}

INNESOCONCHA FLAVESCENS (IREDALE, 1944) (FIGS 5F, 6D, E, 16D–F, 18, 19), Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 61, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334."]}

INNESOCONCHA IREDALE, 1944 Innesoconcha Iredale, 1944: 325. Type species Helix (Microcystis) catletti Brazier, 1872 (by original designation). Melloconcha Iredale, 1944: 323. Type species Melloconcha delecta Iredale, 1944 (by original designation), synon. nov. Tribocystis Iredale, 1944: 323. Type species Tribocystis rosacea Iredale, 1944 (by original designation), synon. nov. Annacharis Iredale, 1944: 324. Type species Annacharis miranda Iredale, 1944 (by original designation), synon. nov. Synonymy of Tribocystis and Annacharis with Melloconcha follows Smith (1992). Description External morphology: Shell small to medium-sized [shell width(hereafter,SW) 3.7–12.0mm, shellheight (hereafter, SH) 1.8–7.8 mm], 4.5–5.5 whorls, glossy, pale golden to dark amber-brown, sometimes with a darker peripheral band. Shell shape discoidal, trochoidal or domed with a low to elevated spire; whorl profile flattened or rounded above and below a rounded to angulate periphery; sutures weakly impressed to strongly channelled. Protoconch sculptured with incised spiral grooves; teleoconch smooth or sculptured with microscopic spiral grooves. Aperture ovately lunate, width greater than height, lip thin; columella smooth or with a small tooth. Internal walls of early whorls complete; umbilicus closed. Right shell lappet long, narrow, finger-shaped; left shell lappet narrow, finger-shaped, sometimes reduced or absent. Caudal horn moderately small to moderately large; caudal foss a horizontal slit in tail., Published as part of Hyman, Isabel T., Caiza, Jennifer & Köhler, Frank, 2023, Dissecting an island radiation: systematic revision of endemic land snails on Lord Howe Island (Gastropoda: Stylommatophora: Microcystidae), pp. 20-75 in Zoological Journal of the Linnean Society 197 (1) on page 49, DOI: 10.1093/zoolinnean/zlac075, http://zenodo.org/record/7499138, {"references":["Iredale T. 1944. The land Mollusca of Lord Howe Island. Australian Zoologist 10: 299 - 334.","Brazier J. 1872. Descriptions of six new species of land shells from Australia and Lord Howe's Island. Proceedings of the Zoological Society of London 1872: 617 - 619.","Smith BJ. 1992. Non-marine Mollusca. In: Houston WWK, ed. Zoological catalogue of Australia, Vol. 8. Canberra: Australian Government Publishing Service, 1 - 408."]}

Malandena Iredale, 1937 Malandena Iredale, 1933: 56 [nom. nud.]. Malandena Iredale, 1937c: 7. Type species Macrochlamys suturalis Odhner, 1917 (OD). Description. External morphology: Shell of about 4.8 whorls, glossy, spire and apex slightly raised. Protoconch smooth; teleoconch sculptured with spiral grooves that become slightly raised spiral lirae. Whorl profile flattened above and rounded below evenly convex periphery. Internal walls of early whorls complete; umbilicus open, very narrow U-shape. Mantle laps of medium length, narrow, tapering to a fine point, not fused, uniform in colour. Right and left mantle lobes small, median lobe absent. Sole of foot and caudal apparatus as for family. Mantle cavity: As for family. Mantle with faintly visible minor blood vessels, pigmentation of white spots. Digestive system: Small oesophageal crop present. Genital system: As for family. Carrefour embedded in albumen gland, talon free. Free oviduct short; capsular gland absent; oviduct internally with irregular longitudinal pilasters. Bursa copulatrix inserted on vagina, moderately long, more than half spermoviduct length; duct of bursa copulatrix wide, distinguishable from bursa copulatrix, internally with longitudinal pilasters. Vagina internally with longitudinal pilasters. Epiphallus enters penis through simple pore; penis internally smooth, longitudinal penis pilasters present, penial diverticulum absent. Penial sheath present, enclosing only penis; penis retractor muscle attached to epiphallus. Epiphallus internally with longitudinal pilasters. Epiphallic retractor caecum absent. Flagellum present; flagellum and distal part of epiphallus without internal cryptae and externally smooth and slender. Spermatophore not present in dissected material. Radula: Relatively long and narrow. Central tooth with small ectocones; mesocone lanceolate, shorter than tooth base. Lateral and marginal tooth fields distinguishable. Lateral teeth with endocone absent; ectocone equal in size to those on central tooth; mesocone shorter than tooth base. Marginal teeth with endocones absent; ectocones approximately same length and breadth as mesocone, not subdivided into extra teeth. Remarks. This genus was introduced by Iredale (1933, 1937c) for Macrochlamys suturalis Odhner, 1917 and placed in the family Macrochlamydidae (following Odhner 1917) due to its large caudal horn and mantle lobes. The group was moved to Sesarinae by Burch (1976), along with Nitor. The genitalia, radula and jaw of Macrochlamys suturalis were described and figured by Odhner (1917). Malandena can be differentiated from all other helicarionid snails with a slender flagellum by its lack of an epiphallic caecum., Published as part of Hyman, Isabel T. & Ponder, Winston F., 2010, A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462, pp. 1-148 in Zootaxa 2462 (1) on page 67, DOI: 10.11646/zootaxa.2462.1.1, http://zenodo.org/record/5312710, {"references":["Iredale, T. (1933) Systematic notes on Australian land shells. Records of the Australian Museum, 19, 37 - 59.","Iredale, T. (1937 c) A basic list of the land Mollusca of Australia - part II. Australian Zoologist, 9, 1 - 39.","Odhner, N. H. J. (1917) Results of Dr. E. Mjobergs Swedish Scientific Expeditions to Australia 1910 - 1913. XVII. Mollusca. Kungliga Svenska Vetenskapsakademiens Handlingar., 52, 1 - 115.","Burch, J. B. (1976) Outline of classification of Australian terrestrial molluscs (native and introduced). Journal of the Malacological Society of Australia, 3, 127 - 156."]}

Liardetia Gude, 1913 Type species Helix clayi Liardet, 1876 (OD) [= Helix striolata Pease 1860 non C. Pfeiffer, 1828]. Fiji. Taxonomic decision for subgeneric arrangement: Baker (1941a)., Published as part of Hyman, Isabel T. & Ponder, Winston F., 2010, A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462, pp. 1-148 in Zootaxa 2462 (1) on page 113, DOI: 10.11646/zootaxa.2462.1.1, http://zenodo.org/record/5312710, {"references":["Baker, H. B. (1941 a) Zonitid snails from Pacific Islands Part 3: Genera other than Microcystinae. Bernice P. Bishop Museum Bulletin, 166, 205 - 346."]}

Stanisicarion n. gen. Type species Helixarion freycineti Férussac, 1821. Etymology. This genus is named for John Stanisic, both in thanks for helpful discussion and assistance over the course of this project and in acknowledgement of his extensive work on land snails in Australia, including Helicarionidae. Description. External morphology: Shell reduced, 3–3.2 whorls, glossy, spire and apex flattened (Stanisicarion freycineti) to slightly raised (S. virens). Protoconch sculptured with incised spiral grooves; teleoconch with microscopic spiral grooves. Whorl profile rounded above and below evenly convex periphery. Internal walls of early whorls complete; umbilicus closed. Mantle laps wide at base, rounded, tapering to blunt point, not fused. Mantle laps uniform in colour. Mantle lobes large to very large, left and median lobes fused to form small lobe over head. Sole of foot and caudal apparatus as for family; caudal foss vertical slit in tail. Mantle cavity: As for family. Mantle with visible minor blood vessels; pigmentation of black spots (not examined in S. virens). Digestive system: Oesophageal crop present. Genital system: As for family. Carrefour embedded in albumen gland; talon embedded (S. virens) or free (S. freycineti). Free oviduct of medium length; capsular gland present; internal sculpture absent (S. virens) or longitudinal pilasters present (S. freycineti). Bursa copulatrix inserted on vagina, short, less than half spermoviduct length; duct of bursa copulatrix wide, distinguishable from bursa copulatrix, internally smooth. Vagina internally with longitudinal pilasters. Epiphallus enters penis through short verge; penis internally covered in pustules, longitudinal penis pilasters absent, diverticulum on penis absent. Penial sheath enclosing only base of penis; penis retractor muscle attached to tip (S. freycineti) or base (S. virens) of epiphallic caecum. Epiphallus internally with longitudinal pilasters. Epiphallic retractor caecum moderate in size, positioned in middle of epiphallus. Flagellum present; flagellum and distal part of epiphallus without internal cryptae and externally smooth and slender. Spermatophore not present in dissected specimens. Radula (based only on freycineti): Relatively long and narrow. Central tooth with small ectocones; mesocone lanceolate, longer than tooth base. Lateral and marginal tooth fields distinguishable. Lateral teeth with endocone smaller than central tooth ectocones; ectocone equal in size to those on central tooth; mesocone longer than tooth base. Marginal teeth with endocones absent; ectocones shorter and narrower than mesocone, not subdivided into extra teeth. Remarks. Stanisicarion can be distinguished from Eungarion, Thularion and Parmacochlea by its less reduced shell, and from Fastosarion by its smaller size and smaller mantle laps and lobes. Anatomically, Stanisicarion differs from the other genera in this section by the presence of a short penial sheath covering only the bottom half of the penis, and by the presence of pustules and absence of pilasters in the penis interior., Published as part of Hyman, Isabel T. & Ponder, Winston F., 2010, A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462, pp. 1-148 in Zootaxa 2462 (1) on page 49, DOI: 10.11646/zootaxa.2462.1.1, http://zenodo.org/record/5312710, {"references":["Ferussac, A. E. J. P. J. F. d'Audebard de (1821) Tableaux systematiques des animaux mollusques classes en familles naturelles, dans lesquels on a etabli la concordance de tous les systemes; suivis d'un Prodrome general pour tous les mollusques terrestres ou fluviatiles, vivants ou fossiles. A. Bertrand, Paris, 184 pp."]}