Helix inconspicua Forbes, 1851: 379 Helix inconspicua C. B. Adams, 1849 Helix rustica Pfeiffer, 1852: 112 Helix inconspicua Forbes, 1851 Helix impexa Reeve, 1852 Helix inconspicua Forbes, 1851 Helix villaris Reeve, 1854 Helix crotali Cox, 1864: 13 Helix inconspicua Forbes, 1851 Helix ( Xesta ) rustica : Cox, 1868: 2 Helix rustica : Brazier, 1876b: 117–118 Helix villaris : Brazier, 1876b: 118 Nanina ( Macrochlamys ) villaris : Pfeiffer & Clessin, 1881: 44 Nanina rustica : Pfeiffer & Clessin, 1881: 46 Nanina impexa : Pfeiffer & Clessin, 1881: 46 Charopa rustica : Tryon, 1886: 212 Nanina Macrochlamys villaris : Tryon, 1886 Charopa rustica : Cox, 1909: 8 Microcystis rustica : Hedley, 1912: 260 Odhner, 1917: 77 Expocystis rustica : Iredale, 1937c: 4 Smith, 1992: 230 A morphological phylogenetic analysis and generic revision of Australian Helicarionidae (Gastropoda: Pulmonata: Stylommatophora), and an assessment of the relationships of the family 2462 Hyman, Isabel T. Ponder, Winston F. Zootaxa 2010 2010-05-14 2462 1 1 148 7RMPQ (Pfeiffer, 1852) Snail Pfeiffer 1852 [151,561,151,177] Gastropoda Helicarionidae Expocystis Animalia Stylommatophora 65 66 Mollusca species rustica   Figures 17P–R, 18H, 19J, 20I, 21J, 22D, 23C      Helix inconspicua Forbes, 1851: 379[ non  Helix inconspicuaC. B. Adams, 1849].     Helix rustica Pfeiffer, 1852: 112[ nom. nov.for  Helix inconspicua Forbes, 1851].   Helix impexa Reeve, 1852: pl. 130, sp. 795 [ nom. nov.for  Helix inconspicua Forbes, 1851].   Helix villaris Reeve, 1854: pl. 195, sp. 1375.     Helix crotali Cox, 1864: 13[ nom. nov.for  Helix inconspicua Forbes, 1851].     Helix( Xesta) rustica: Cox, 1868: 2, pl. 9, figs 3, 3a.    Helix(T halassia)   rustica: Brazier, 1876b: 117–118.    Helix(T halassia)   villaris: Brazier, 1876b: 118.     Nanina( Macrochlamys) villaris: Pfeiffer & Clessin, 1881: 44.    Nanina(T halassia)   rustica: Pfeiffer & Clessin, 1881: 46.    Nanina(T halassia)   impexa: Pfeiffer & Clessin, 1881: 46.    Charopa(T halassia)   rustica: Tryon, 1886: 212, pl. 62, figs 44–45.   Nanina(Subg. Xesta Sect. Macrochlamys)  villaris: Tryon, 1886: pl. 105, pl. 35, fig. 41.    Charopa(T halassia)   rustica: Cox, 1909: 8.     Microcystis rustica: Hedley, 1912: 260, pl. viii, fig. 42;  Odhner, 1917: 77, figs 29, 30c, 33.     Expocystis rustica: Iredale, 1937c: 4;  Smith, 1992: 230. Synonymy follows Iredale (1937c: 4)and Smith (1992: 230).   Material examined.   Queensland, Australia: BMNH 1982243(  H. villaris holotype) (one shell measured, photographed), Lizard Island;  AM C425540( two specimensdissected, one radula examined), S of Cooktown, Mt. Cook National Park( 15º30' S, 145º15' E), on ground, semi deciduous microphyll vine forest,  23 Oct. 1975, P.H. Colman.   Description. External morphology: Shell ( Figures 19J, 20L, 21J) of about 5 whorls, very light pinkishbrown; shape and sculpture as for genus. Animal white. Mantle laps and lobes as for genus. Caudal horn large; caudal foss vertical slit in tail.  Mantle cavity and digestive system: As for genus.  Genital system( Figures 18H, 22D, 23C): As for genus. Penis moderately long; epiphallus enters penis through simple pore; penis internally covered in irregular ridges, four longitudinal penis pilasters present. Epiphallus longer than penis. Spermatophore soft capsule with firm tail pipe open at one end, tail pipe long, sculptured with longitudinal rows of tiny teeth.   Radula( Figure 17P–R): As for genus. Radular formula (30.14.1.14.30) × 100 rows.  Range and habitat.  Expocystis rusticais found from Innisfail to Cape YorkPeninsula, N Queensland, on the mainland and also on many islands of the Great Barrier Reef. It is found in rainforest, microphyll vine forest and under trees on or behind beaches, in leaf litter, inside rotting logs and under stones.   Remarks.The synonymy of  Helix inconspicua Forbes, 1851,  H. impexa Reeve, 1852and  H. crotali Cox, 1864with  Expocystis rusticawas recognised by most early authors from Cox (1868)onwards. However, Cox (1868)incorrectly placed  Pravonitor krefftiin synonymy with  Helix villaris. Iredale (1937c)first recognised  H. villarisas a synonym of  Expocystis rusticaand this was followed by Smith (1992)and Smith et al.(2002). The anatomical details given here differ from those presented by Odhner (1917), who illustrated the penial complex. His figure and description indicate that the epiphallus is absent and that the flagellum and penis retractor muscle are both present at the junction of the penis and vas deferens. A dart sac is also described as being present. Some of these differences are, in part at least, due to misinterpretation. The structure referred to as the dart sac is actually the bursa copulatrix (see also remarks under  Malandena suturalisbelow). It is possible that the structure referred to as the flagellum is actually the caecum, although this interpretation assumes that Odhner omitted the actual flagellum from the drawing.  The radulashown here also differs slightly from that figured by Odhner (1917), who shows his specimen as having unicuspid marginal teeth. However, in his written description of the radula, Odhnerstates that the secondary cusp of the marginal teeth is ‘rather reduced in size and rather remote from the tooth end’ ( Odhner 1917: 81), implying that there is a small secondary cusp present. A small secondary cusp was present in the radulaexamined in the current study. It is possible that Odhner’s (1917)specimen was misidentified, but no shell was figured and Odhner’s material was not examined, so this cannot be confirmed. BMNH Australia Lizard Island 65 66 BMNH 1982243 1 Queensland holotype 1975-10-23 AM P. H. Colman Australia -15.5 Cooktown 1284 145.25 Mt. Cook National Park 65 66 C425540 1 Queensland holotype