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FIGURE 38. 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 38. Mantle cavities. A. Everettia sp., BPBM 20020188. B. Coneuplecta pampini, AM C438751. C. Hiona pilsbryi, BPBM 20680 & 20701 (mixed). D. Trochomorpha rubens, FMNH 175247. Scale bars 2 mm (B–D), 5 mm (A). Abbreviations: i, intestine; k, kidney; mc, mantle collar; pc, pericardium; pv, pulmonary vein; ur1, primary ureter; ur2, secondary ureter.
FIGURE 25. 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 25. Radulae. SEM. A–C. Malandena suturalis, AM C425085. D–F Westracystis lissus, AM C205297. G–I. Echonitor albumenoidea, SAM D16567. J–L. Tarocystis responsivus, AM C425327. M–O. New Caledonian species NC3, MNHN no registration number. P–R. New Caledonian species NC5, MNHN no registration number. Scale bars 20 mm (A–R).
FIGURE 37. Spermatophores. SEM. A, B. Elisolimax flavescens, NMSA L5563. C, D 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 37. Spermatophores. SEM. A, B. Elisolimax flavescens, NMSA L5563. C, D. Macrochlamys petrosa, AM C205305. E, F. Orpiella concavum, BMNH no registration number. Scale bars 0.2 mm (D, F), 0.5 mm (B), 1 mm (A, C, E).
FIGURE 33. 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 33. Penis interiors. A. Erepta odontina, AM C205302. B. Kalidos balstoni, AM C205303. C. Madagascan species MA1, AM C205304. D. Sheldonia poeppigii, NMSA L5563. E. Elisolimax flavescens, NMSA V9710. F. Parmarion martensi, BPBM 252738. G. Coneuplecta pampini, AM C438751. H. Hiona pilsbryi, BPBM 206801 & 20701. I. Trochomorpha rubens, FMNH 175247. J. Videna planorbis, FMNH 160957. K. Dendrotrochus eva eva, FMNH 175196. L. Orpiella concavum, BMNH no registration number. Scale bars 1 mm (A, C–D, F–H, J, M), 2 mm (B, E, I, K). Abbreviations: bc, bursa copulatrix; c, caecum; ep, epiphallus; fov, free oviduct; ovi, ovipositer; p, penis; pp, penial pilaster; prm, penis retractor muscle; ppu, penial pustules; pv, penial verge.
FIGURE 28. Shells, ventral view. A. Westracystis lissus, BMNH90.12.30.80–4 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 28. Shells, ventral view. A. Westracystis lissus, BMNH90.12.30.80–4 (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.
FIGURE 35 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 35. External morphology of caudal regions. A. Kalidos balstoni, AM C205303. B. Macrochlamys petrosa, AM C205303. C. Everettia sp., BMNH 20020188. D. Greenwoodoconcha nux, AM C433375. E. Videna planorbis, FMNH 160957. F. Dendrotrochus eva eva, FMNH 160957. Scale bars 1 mm (D), 2 mm (E–F), 5 mm (A–C).
FIGURE 24. 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 24. Reproductive systems. A. Malandena suturalis, AM C425085. B. Westracystis lissus, FMNH 211717. C. Echonitor cyrtochila, SAM D16557. D. Echonitor albumenoidea, SAM D16567. E. Periclocystis ardeni, SAM D16562. F. Tarocystis responsivus, AM C425327. G. New Caledonian species NC3, MNHN no registration number. H. New Caledonian species NC5, MNHN no registration number. Scale bars 1 mm (C, G), 2 mm (A–B, D–F, H). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; m, muscle attaching penial sheath to epiphallus (only shown in D & G); ov, ovotestis; p, penis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; t, talon; ut, uterus; v, vagina; vd, vas deferens.
FIGURE 18. 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 18. Reproductive systems. A. Stanisicarion freycineti, AM C391188. B. Stanisicarion virens, QMMO19238. C. Eungarion mcdonaldi, AM C438760. D. Parmacochlea fischeri, AM C441743. E. Nitor circumcincta, AM C356607. F. Nitor subrugata, AM C334537. G. Pravonitor kreffti, AM C425415. H. Expocystis rustica, AM C425540. Scale bars 1 mm (E), 2 mm (A–D, G–H), 5 mm (F). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; m, muscle attaching penial sheath to epiphallus (only shown in A); ov, ovotestis; p, penis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; ut, uterus; v, vagina; vd, vas deferens.
FIGURE 12. 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 12. Reproductive systems. 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 C348670. H. Brevisentis jacksoniensis, AM C205293. I. Fastosarion superba, AM C425248. Scale bars 2 mm (C–D, F–H), 5 mm (A–B, E, I). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; m, muscle attaching penial sheath to epiphallus (only shown in E); ov, ovotestis; p, penis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; ut, uterus; v, vagina; vd, vas deferens.
FIGURE 15. 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 15. Radulae. SEM. A–C Helicarion mastersi, AM C334204. D–F. Mysticarion porrectus, QMMO29298. G–I. Mysticarion hyalina, AM C128630. J–L. Parmavitrina planilabris, AM C334357. M–O. Peloparion helenae, AM C205295. P–R. Brevisentis jacksoniensis, AM C205293. Scale bars 20 mm (A–I, M–R); 50 mm (J–L).
FIGURE 21. 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 21. Shells, ventral view. A. Stanisicarion freycineti, AM C391188. B. Stanisicarion virens, BMNH 197776 (S). C. Eungarion mcdonaldi, QMMO35628 (H). D. Parmacochlea fischeri, AM C432179. E. Nitor circumcincta, AM C104161 (H). F. Nitor subrugata, AM C333711. G. Nitor subrugata, AM C63479 (S of N. subrugata helmsianus). H. Pravonitor kreffti, BMNH no number (POS). I. Pravonitor kreffti, AM C29647 (H of P. kreffti insularum). J. Expocystis rustica, BMNH1982243 (POH of Helix villaris). K. Malandena suturalis, AM C425087. Scale bars 3 mm (A, E, J), 5 mm (B–D, F–I, K). Photographs by H. Barlow (A, D, F, I), D. Beechey (E, G), I. Hyman (K), W. Ponder (B, H, J), copyright Australian Museum; photographer unknown, copyright Queensland Museum (C). Abbreviations: H, holotype; POH, possible holotype; POS, possible syntype; S, syntype.
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.
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.
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).
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.
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).
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.
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.
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.
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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