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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."]}