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FIGURE 27. Shells, lateral view. A. Westracystis lissus, BMNH 90.12.30.80–4 in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 27. Shells, lateral view. A. Westracystis lissus, BMNH 90.12.30.80–4 (S). B. Westracystis lissus, AM C64901 (S of W. tentus). C. Echonitor cyrtochila, BMNH 1922.8.29.16 (H). D. Echonitor albumenoidea, AM C60533 (H). E. Periclocystis ardeni, AM C60541 (H). F. Tarocystis responsivus, AM C104308 (S). G. Conueplecta (Durgellina) calculosa, AM C374873. H. Coneuplecta (Nevelasta) pampini, AM C126763. Scale bars 2 mm (B, G–H), 3 mm (A, C–F). Photographs by H. Barlow (B, F–H), D. Beechey (D–E), W. Ponder (A, C). Abbreviations: H, holotype; S, syntype.

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FIGURE 9. Shells, ventral view. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 9. Shells, ventral view. A. Helicarion cuvieri, AM C443929. B. Helicarion cuvieri, BMNH 197780 (S of Vitrina verrauxii). C. Helicarion mastersi, AM C101139 (H). D. Helicarion mastersi, AM C101138 (PRH of H. mastersi callidus). E. Mysticarion insuetus, AM C113776 (S). F. Mysticarion porrectus, AM C101136 (PRH). G. Mysticarion hyalina, BMNH 1983075 (H). H. Parmavitrina planilabris, AM C103630. I. Peloparion helenae, AM C101142 (H). J. Brevisentis jacksoniensis, BMNH not registered (S). K. Brevisentis jacksoniensis, AM C101143 (H of B. exclusus). L. Fastosarion superba, BMNH not registered (POS). Scale bars 3 mm (D–G, I–K), 5 mm (A–C, H), 10 mm (L). Photographs by H. Barlow (A, C–D, H–I), D. Beechey (F, K), W. Ponder (B, E, G, J, L). Abbreviations: H, holotype; POS, possible syntype; PRH, probable holotype; S, syntype.

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FIGURE 14. Spermatophores. SEM. A, B in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 14. Spermatophores. SEM. A, B. Helicarion cuvieri, AM C424333. C. Helicarion mastersi, AM C334204. D. Brevisentis jacksoniensis, AM C205293. E, F. Eungarion mcdonaldi, AM C438760. G, H. Nitor wiangariensis, AM C334361. Scale bars 0.1 mm (F), 0.2 mm (H), 0.5 mm (B, C, G), 1 mm (A, D, E).

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FIGURE 7. Shells, dorsal view. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 7. Shells, dorsal view. A. Helicarion cuvieri, AM C443929. B. Helicarion cuvieri, BMNH 197780 (S of Vitrina verrauxii). C. Helicarion mastersi, AM C101139 (H). D. Helicarion mastersi, AM C101138 (PRH of H. mastersi callidus). E. Mysticarion insuetus, AM C113776 (S). F. Mysticarion porrectus, AM C101136 (PRH). G. Mysticarion hyalina, BMNH 1983075 (H). H. Parmavitrina planilabris, AM C103630. I. Peloparion helenae, AM C101142 (H). J. Brevisentis jacksoniensis, BMNH not registered (S). K. Brevisentis jacksoniensis, AM C101143 (H of B. exclusus). L. Fastosarion superba, BMNH not registered (POS). Scale bars 3 mm (G, I–K), 5 mm (A–F, H), 10 mm (L). Photographs by M. Allan (D, I), H. Barlow (A, C, H), D. Beechey (E–F, K), W. Ponder (B, G, J, L). Abbreviations: H, holotype; POS, possible syntype; PRH, probable holotype; S, syntype.

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FIGURE 6. Live Helicarionidae. A–B in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 6. Live Helicarionidae. A–B. Helicarion mastersi, AM C399075. Whole animal (A); close-up of caudal region (B) (arrow shows caudal mucous plug). C. Eggs of Mysticarion porrectus, AM C421841. D. Parmavitrina megastoma, AM C377461. E. Parmavitrina sp., AM C398947. F. Peloparion iridis, AM C161946. G. Peloparion iridis, Mt. Ravensbourne. H–I. Fastosarion superba. J. Stanisicarion freycineti, AM C391188. K. Stanisicarion virens. L. Eungarion mcdonaldi. M. Eggs of Eungarion mcdonaldi. N. Parmacochlea fischeri, AM C106767. Photographs by A. Miller (A–C, E, J), M. Shea (D), W. Rudman (F), F. Millen (N), all copyright Australian Museum; photographer unknown, copyright Queensland Museum (G–I, K–M).

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FIGURE 4 in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 4. Australian Helicarionidae. Majority-rule consensus tree of 13,839 trees, length 99. Where two or more species from a genus are included, the genera are shaded. Groups A and B are referred to in text.

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FIGURE 10. Mantle cavities. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 10. Mantle cavities. A. Helicarion cuvieri, AM C424333. B. Brevisentis jacksoniensis, AM C356645. C. Stanisicarion freycineti, AM C391188. D. Nitor circumcincta, AM C365822. E. Tarocystis responsivus, AM C425327. F. New Caledonian species NC3, MNHN no registration number. Scale bars 1 mm (D–E), 2 mm (A–C, F). Abbreviations: dg, digestive gland; i, intestine; k, kidney; mc, mantle collar; mlo, median mantle lobe; pc, pericardium; pn, pneumostome; pv, pulmonary vein; rla, right mantle lap; rlo, right mantle lobe; ur1, primary ureter; ur2, secondary ureter.

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FIGURE 26. Shells, dorsal view. A. Westracystis lissus, BMNH 90.12.30.80–4 in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 26. Shells, dorsal view. A. Westracystis lissus, BMNH 90.12.30.80–4 (S). B. Westracystis lissus, AM C64901 (S of W. tentus). C. Echonitor cyrtochila, BMNH 1922.8.29.16 (H). D. Echonitor albumenoidea, AM C60533 (H). E. Periclocystis ardeni, AM C60541 (H). F. Tarocystis responsivus, AM C104308 (S). G. Conueplecta (Durgellina) calculosa, AM C374873. H. Coneuplecta (Nevelasta) pampini, AM C126763. Scale bars 2 mm (B, G–H), 3 mm (A, C–F). Photographs by M. Allan (F), H. Barlow (B, G–H), D. Beechey (D–E), W. Ponder (A, C). Abbreviations: H, holotype; S, syntype.

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FIGURE 3 in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 3. Majority-rule consensus tree (770 trees, length 117 steps) resulting from the morphological analysis with two additional mantle lobe characters. Families and subfamilies are shaded and are labelled in bold if the group is not monophyletic.

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FIGURE 2 in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 2. Majority-rule consensus tree (646 trees, length 106 steps) from the morphological analysis with Dyakiidae and Urocyclidae excluded. Families and subfamilies are shaded and are labelled in bold if the group is not monophyletic.

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FIGURE 36. Radulae. SEM. A–C. Sheldonia poeppigii, NMSA V9710. D–F. Elisolimax flavescens, NMSA L5563. G– I in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 36. Radulae. SEM. A–C. Sheldonia poeppigii, NMSA V9710. D–F. Elisolimax flavescens, NMSA L5563. G– I. Macrochlamys petrosa, AM C205305. J–L. Sesara infrendens, BMNH 1995067. M–O. Parmarion martensi, BPBM 252738. P–R. Everettia sp., BMNH 20020188. Scale bars 10 mm (G–I), 20 mm (A–F, J–R).

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FIGURE 13. Penis interiors. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 13. Penis interiors. A. Helicarion cuvieri, AM C424333. B. Helicarion mastersi, AM C334204. C. Mysticarion porrectus, AM C205296. D. Mysticarion hyalina, AM C128630. E. Parmavitrina planilabris, AM C334357. F. Peloparion helenae, AM C205295. G. Brevisentis jacksoniensis, AM C205293. H. Fastosarion superba, AM C425248. I. Stanisicarion freycineti, AM C391188. J. Stanisicarion virens, QMMO19238. K. Eungarion mcdonaldi, AM C438760. L. Parmacochlea fischeri, AM C441743. Scale bars 1 mm (A–D, F–G I, K–L), 2 mm (E, H, J). Abbreviations: c, caecum; ep, epiphallus; fl, flagellum; p, penis; pp, penial pilaster; ppu, penial pustules; prm, penis retractor muscle; pt, penial sheath; pv, penial verge; vd, vas deferens.

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FIGURE 8. Shells, lateral view. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 8. Shells, lateral view. A. Helicarion cuvieri, AM C443929. B. Helicarion cuvieri, BMNH 197780 (S of Vitrina verrauxii). C. Helicarion mastersi, AM C101139 (H). D. Helicarion mastersi, AM C101138 (PRH of H. mastersi callidus). E. Mysticarion insuetus, AM C113776 (S). F. Mysticarion porrectus, AM C101136 (PRH). G. Mysticarion hyalina, BMNH 1983075 (H). H. Parmavitrina planilabris, AM C103630. I. Peloparion helenae, AM C101142 (H). J. Brevisentis jacksoniensis, BMNH not registered (S). K. Brevisentis jacksoniensis, AM C101143 (H of B. exclusus). L. Fastosarion superba, BMNH not registered (POS). Scale bars 3 mm (E–G, I–K), 5 mm (A–D, H), 10 mm (L). Photographs by H. Barlow (A, C–D, G–H, L), D. Beechey (E, J), W. Ponder (B, F, I, K). Abbreviations: H, holotype; POS, possible syntype; PRH, probable holotype; S, syntype.

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FIGURE 30. Whole radulae. A. Peloparion iridis, QMMO51026. B in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 30. Whole radulae. A. Peloparion iridis, QMMO51026. B. Caldwellia imperfecta, AM C205300. C. Coneuplecta pampini, AM C438751. D. Cystopelta astra, AM C354840. E. Parmellops sp., AM C393245. Scale bars 0.5 mm (A–C), 1 mm (D–E).

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FIGURE 29. Reproductive systems. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 29. Reproductive systems. A. Caldwellia angularis, AM C205299. B. Caldwellia imperfecta, AM C205300. C. Erepta odontina, AM C205302. D. Ctenophila caldwelli, AM C205301. E. Kalidos balstoni, AM C205303. Scale bars 1 mm (E), 2 mm (B, D), 5 mm (A, C). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ca, carrefour; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; ov, ovotestis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; ut, uterus; v, vagina; vd, vas deferens.

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FIGURE 32. Mantle cavities. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 32. Mantle cavities. A. Caldwellia imperfecta, AM C205300. B. Madagascan species MA1, AM C205304. C. Sheldonia poeppigii, NMSA V9710. D. Macrochlamys petrosa, AM C205303. E. Parmarion martensi, BPBM 252738. Scale bars 2 mm (A–B, E), 3 mm (C), 5 mm (D). Abbreviations: dg, digestive gland; i, intestine; k, kidney; mc, mantle collar; pc, pericardium; pn, pneumostome; pv, pulmonary vein; ur1, primary ureter; ur2, secondary ureter.

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FIGURE 34. Reproductive systems. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 34. Reproductive systems. A. Madagascan species MA1, AM C205304. B. Elisolimax flavescens, NMSA V9710. C. Sheldonia poeppigii, NMSA L5563. D. Macrochlamys petrosa, AM C205305. E. Sesara infrendens, BMNH 1995067. F. Parmarion martensi, BPBM 252738. Scale bars 1 mm (E), 2 mm (A), 5 mm (B–D, F). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; ls, lime-sac; m, muscle attaching penial sheath to epiphallus (only shown in E); ov, ovotestis; p, penis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; st, stimulator; ut, uterus; v, vagina; vd, vas deferens.

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FIGURE 23. Spermatophores. SEM. A, B in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 23. Spermatophores. SEM. A, B. Nitor subrugata, AM C334537. C. Expocystis rustica, AM C425540. D. Ctenophila caldwelli, AM C205301. E, F. New Caledonian species NC5, MNHN no registration number. G, H. Kalidos balstoni, AM C205303. Scale bars 0.2 mm (B, F, H), 0.5 mm (C, E), 1 mm (A, D, G).

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FIGURE 17. Radulae. SEM. A–C in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 17. Radulae. SEM. A–C. Fastosarion superba, AM C425248. D–F. Stanisicarion freycineti, AM C391188. G– I. Parmacochlea fischeri, AM C441743. J–L. Nitor circumcincta, AM C356607. M–O. Pravonitor kreffti, AM C425415. P–R. Expocystis rustica, AM C425540. Scale bars 20 mm (D–R), 50 mm (A–C).

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FIGURE 22. Penis interiors. A in A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462

FIGURE 22. Penis interiors. A. Nitor circumcincta, AM C356607. B. Nitor subrugata, AM C334537. C. Pravonitor kreffti, AM C425415. D. Expocystis rustica, AM C425540. E. Malandena suturalis, AM C425085. F. Echonitor cyrtochila, SAM D16557. G. Echonitor albumenoidea, SAM D16567. H. Periclocystis ardeni, SAM D16562. I. New Caledonian NC3, MNHN no registration number. J. New Caledonian NC5, MNHN no registration number. K. Caldwellia angularis, AM C205299. L. Caldwellia imperfecta, AM C205300. Scale bars 1 mm (A–L). Abbreviations: ep, epiphallus; pd, penial diverticulum; pp, penial pilaster; ppd, diamond-shaped, ridged pustules; prm, penis retractor muscle; ppu, penial pustules; pv, penial verge.

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