Sheldonia poeppigii ( Pfeiffer, 1846 )

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 : 93-94

Sheldonia poeppigii ( Pfeiffer, 1846)

Figures 32C , 33D , 34C , 36A–C

Material examined. South Africa: NMSA V9710 (one specimen dissected, radula examined), Pietermaritzburg area , Umgeni River Valley , above Table Mountain , Cumberland Conservancy, Kwa Zulu-Natal (29º30.129' S, 30º30.52' E), in leaf litter under bush clumps in open valley bushveld, 19 Dec. 2001, D. Herbert, L. Davis & L. Packer .

Description. External morphology: Shell 3.5 whorls, golden brown, glossy, spire and apex slightly raised. Protoconch sculptured with no sculpture; teleoconch with microscopic spiral grooves. Whorl profile rounded above and below evenly convex periphery. Internal walls of early whorls complete; umbilicus closed. Animal white and grey. Right mantle lap absent, left mantle lap tiny, narrow, finger-shaped. Right mantle lobe small, left and median mantle lobes very small. Sole of foot and caudal apparatus as for family; caudal horn very long, overhanging foot; caudal foss vertical slit in tail.

Mantle cavity ( Figure 32C ): As for family. Mantle with visible minor blood vessels, pigmentation of white spots.

Digestive system: Oesophageal crop absent.

Genital system ( Figures 33D , 34C ): As for family. Carrefour and talon both not embedded in albumen gland. Free oviduct long to very long; capsular gland present; internal longitudinal pilasters present. Bursa copulatrix inserted on vagina, short, less than half spermoviduct length; duct of bursa copulatrix narrow, distinguishable from bursa copulatrix, internally with longitudinal pilasters. Vagina internally with longitudinal pilasters. Stimulator absent. Penis long; epiphallus enters penis through simple pore; penis internally covered in pustules, ten longitudinal penis pilasters present, penial diverticulum absent. Penial sheath present, enclosing only penis; penis retractor muscle attached to junction of penis and epiphallus. Epiphallus shorter than penis, internally with longitudinal pilasters. Epiphallic retractor caecum very long, positioned at penisepiphallus junction. Flagellum present; flagellum and distal part of epiphallus without internal cryptae and externally smooth and slender. Spermatophore not present in dissected specimens.

Radula ( Figure 36A–C ): Relatively long and narrow. Central tooth with small ectocones; mesocone lanceolate, longer than tooth base. Lateral and marginal tooth fields distinguishable. Lateral teeth with endocone smaller than central tooth ectocones; ectocone equal in size to those on central tooth; mesocone slightly shorter than tooth base. Marginal teeth with endocones absent; ectocones shorter and narrower than mesocone, not subdivided into extra teeth. Radular formula (72.13.1.13.72) × 102 rows.

Remarks. There are no available anatomical data for the type species of Sheldonia , S. trotteriana (Benson, 1848) , so placement of S. poeppigii in this genus cannot be confirmed. Sheldonia is found in South Africa and has at least 30 species and subspecies ( Schileyko 2002b). It is usually included in Urocyclidae as part of Trochonanininae ( Baker 1941a; Hausdorf 1998; Solem 1966); however, Schileyko (2002b) included Sheldonia in Ariophantidae , in the subfamily Trochozonitinae and tribe Sheldoniini . The molecular phylogeny of Hyman et al. (2007) showed S. poeppigii to be the sister group to ( Ariophantidae + Helicarionidae ). In the current study S. poeppigii grouped with Elisolimax flavescens ( Urocyclinae , Urocyclidae ) ( Figure 1 ).

Figures

FIGURE 1. Majority-rule consensus tree (1,743 trees, length 117 steps) from the morphological analysis containing representatives of all groups. Families and subfamilies are shaded and are labelled in bold if the group is not monophyletic.

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.

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 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.

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).