Nitor subrugata ( Reeve, 1852 )

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, Zootaxa 2462 (1), pp. 1-148 : 61-62

Nitor subrugata ( Reeve, 1852)

Figures 18F , 19F–G , 20E–F , 21F–G , 22B , 23A–B

Helix subrugata Reeve, 1852 : pl. 128, sp. 773.

Nanina (T halassia) subrugata: Albers & Martens, 1860: 59 ; Pfeiffer & Clessin, 1881: 47.

? Helix graftonensis Cox, 1864 [page not numbered].

? Helix clarencensis Cox, 1865: 594 .

Helix (Rhyssota) subrugata: Cox, 1868: 4 , pl. 9, figs 2, 2a.

Microcystis marmorata: Odhner, 1917: 78 , figs 30a, 31.

Nitor subrugatus: Iredale, 1937c: 2 ; Iredale, 1941: 3, fig. 7.

Nitor subrugata helmsianus Iredale, 1941: 3 .

Nitor subrugata: Smith, 1992: 239 .

Synonymy follows Iredale (1937c); Smith (1992).

Material examined. New South Wales, Australia: Syntypes: BMNH 1982236 (one shell measured), Australia .

Other material: AM C101140 ( N. subrugata helmsianus syntypes) (one shell measured, photographed), E of Lismore , Wollongbar (28º49' S, 153º24' E), pre 1941 , R. Helms ; AM C333711 (one shell measured, photographed), Grafton , Clarence River (29º41' S, 152º57' E), pre 1912 ; AM C334537 (one specimen dissected), N of Lismore , Whian Whian State Forest, Red Scrub Flora Reserve (28º38' S, 153º20' E), complex notophyll vine forest, 15 May 1976, I. Loch, P.H. Colman .

Description. External morphology: Shell ( Figures 19F–G , 20E–F , 21F–G ) 5.3 whorls, light yellowbrown; whorl profile flattened above and rounded below angulate periphery; umbilicus open, very narrow Ushape; otherwise shell shape and sculpture as for genus. Animal yellow. Right mantle lap very short, narrow, finger-shaped; left mantle lap absent; mantle lobes as for genus. Caudal horn moderately large; caudal foss vertical slit in tail.

Mantle cavity: As for genus. Mantle with no pigmentation.

Digestive system: As for genus.

Genital system ( Figures 18F , 22B , 23A–B ): As for genus. Bursa copulatrix internally smooth. Epiphallus enters penis through short verge; penis internally smooth, four to five longitudinal penis pilasters present. Penial sheath present, enclosing only penis. Epiphallus longer than penis, internally with longitudinal pilasters. Spermatophore as for genus.

Radula: Not examined.

Range and habitat. Nitor subrugata ranges from around Grafton in northern New South Wales to Mt. Tamborine, Queensland, although some specimens are known from further south, around Dungog (AM C374283, AM C 374285). It has been found in notophyll and microphyll vine forest, rainforest, wet sclerophyll forest and scrubs, under logs and rocks and in leaf litter.

Remarks. Nitor helmsianus and N. graftonensis (= clarencensis ) have been synonymised with N. subrugata , following Smith (1992). Nitor helmsianus was originally described as a subspecies of N. subrugata based on its smaller size, higher spire, darker colour with less marked plications and narrower umbilicus. Examination of shells from throughout the range of N. subrugata and measurements of the type material of N. helmsianus and N. subrugata show little difference in size and shape, so Smith’s (1992) synonymy of the two species is followed here.

Nitor graftonensis and N. clarencensis were described by Cox (1864, 1865) in consecutive years. Neither species was illustrated, but both had identical descriptions with the only difference being the specific name used. While it is probable that the two names are synonyms, the types of both species have been lost, which in the absence of figures makes it impossible to accurately determine their status. However, because the dimensions given (diameter 12 mm, height 7 mm, number of whorls 5.5) match those of N. subrugata , the aperture is described as angular and the locality correlates, we treat these names as probable synonyms of N. subrugata .

Figures

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 19. Shells, dorsal view. A. Stanisicarion freycineti, AM C391188. B. Stanisicarion virens, BMNH 197776 (S). C. Eungarion mcdonaldi, QMMO35628. D. Parmacochlea fischeri, AM C432179. E. Nitor circumcincta, AM C104161 (POS of Helix marmorata). 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, BMNH 1982243 (POH of Helix villaris). K. Malandena suturalis, AM C425087. Scale bars 3 mm (E, J), 5 mm (A–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 20. Shells, lateral view. A. Stanisicarion freycineti, AM C391188. B. Stanisicarion virens, BMNH 197776 (S). C. Parmacochlea fischeri, AM C432179. D. Nitor circumcincta, AM C104161 (H). E. Nitor subrugata, AM C333711. F. Nitor subrugata, AM C63479 (S of N. subrugata helmsianus). G. Pravonitor kreffti, BMNH no number (POS). H. Pravonitor kreffti, AM C29647 (H of P. kreffti insularum). I. Expocystis rustica, BMNH 1982243 (POH of Helix villaris). J. Malandena suturalis, AM C425087. Scale bars 3 mm (A, D, I), 5 mm (B–C, E–H, J). Photographs by H. Barlow (A, C, E, H), D. Beechey (D, F), W. Ponder (B, G, I), I. Hyman (J). Abbreviations: H, holotype; POH, possible holotype; POS, possible syntype; S, syntype.

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 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.

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).