Trochomorpha (Hartmanitrochus) rubens Hartman, 1888

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 : 117-118

Trochomorpha (Hartmanitrochus) rubens Hartman, 1888

Figures 11M , 33I , 38D , 41B , 42A–C

Material examined. Vanuatu: FMNH 175247 (two specimens dissected, one radula examined), 3–11 km S of Hog Harbour , plateau top, NE Espiritu Santo, alt. 120 m, 29–30 Oct. 1972, L. Price .

Description. External morphology: Shell of about 6.2 whorls, light orange-brown, glossy, spire and apex raised. Protoconch smooth; teleoconch sculptured with strong radial growth lines. Whorl profile rounded above and below keeled periphery. Internal walls of early whorls complete; umbilicus open, narrow U-shape. Animal light orange-brown. Mantle laps and lobes absent. Sole of foot undivided; caudal apparatus with sole longer than foot ( Muratov’s 1999 zonitid type).

Mantle cavity ( Figure 38D ): As for family. Pigmentation of black and white spots, mantle gland absent.

Digestive system: Oesophageal crop absent.

Genital system ( Figures 33I , 41B ): As for family. Carrefour and talon both not embedded in albumen gland. Free oviduct long; capsular gland present; internal longitudinal pilasters absent. Bursa copulatrix inserted at junction of penis and free oviduct, short, less than half spermoviduct length; duct of bursa copulatrix wide, distinguishable from bursa copulatrix, internally with longitudinal pilasters. Vagina absent. Penis moderately long; epiphallus enters penis through simple pore; penis internally smooth, three major longitudinal penis pilasters present (additional pilasters at base), penial diverticulum absent. Penial sheath absent; penis retractor muscle attached to junction of penis and epiphallus. Epiphallus shorter than penis, internally with longitudinal pilasters. Epiphallic retractor caecum absent; Flagellum and lime-sac absent. Spermatophore not present in dissected specimen.

Radula ( Figure 42A–C ): Relatively long and narrow. Central tooth with no ectocones; mesocone triangular, much longer than tooth base. Lateral and marginal tooth fields distinguishable. Lateral teeth with endocone absent; ectocone much smaller than those on central tooth; mesocone much longer than tooth base. Marginal teeth with endocones absent; ectocones slightly shorter and narrower than the mesocone, not subdivided into extra teeth. Radular formula (33.13.1.13.33) × 109 rows.

Remarks. The anatomy of Trochomorpha rubens is figured by Schileyko (2002a), and agrees with that described above except for the presence of a few additional short pilasters around the base of the penis interior that were seen in the current study but are absent from Schileyko’s (2002a) figure.

The subgeneric arrangement of Schileyko (2002a) is followed in the current study. The subgenus Hartmanitrochus is limited to Vanuatu and the Malaku Islands, and contains two to three species ( Schileyko 2002a).

Trochomorpha rubens grouped with Videna planorbis (Trochomorphidae) in the phylogenetic analysis ( Figures 1–3 ). In the molecular phylogenetic analysis of Wade et al. (2006), the genera Trochomorpha and Videna also grouped together ( Figure 44B ).

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 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.

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.

FIGURE 11. Nervous systems. Cerebral commissure cut (cut regions shaded with diagonal lines). A. Helicarion cuvieri, AM C424333. B. Brevisentis atratus, AM C205294. C. Stanisicarion freycineti, AM C391188. D. Nitor circumcincta, AM C365822. E. Tarocystis responsivus, AM C425327. F. New Caledonian species NC3, MNHN no registration number. G. Ctenophila caldwelli, AM C205301. H. Madagascan species MA1, AM C205304. I. Macrochlamys petrosa, AM C205305. J. Everettia sp., BMNH 20020188. K. Coneuplecta pampini, AM C438751. L. Lamprocystis guttula, FMNH 174940. M. Trochomorpha rubens, FMNH 175247. Scale bars 1 mm except for J (scale bar 2 mm). Abbreviations: ce, cerebral ganglion; pa, parietal ganglion; pe, pedal ganglion; pl, pleural ganglion; vg, visceral ganglion.

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 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 41. Reproductive systems. A. Lamprocystis guttula, FMNH 174940. B. Trochomorpha rubens, FMNH 175247. C. Videna planorbis, FMNH 160957. D. Orpiella concavum, BMNH no registration number. E. Dendrotrochus eva eva, FMNH 175196. Scale bars 1 mm (A), 2 mm (B–D, F), 5 mm (E). Abbreviations: alb, albumen gland; bc, bursa copulatrix; ca, carrefour; e, eggs / embryos; ep, epiphallus; fov, free oviduct; her, hermaphrodite duct; ov, ovotestis; p, penis; pr, prostate; prm, penis retractor muscle; puo, post-uterine oviduct; t, talon; ut, uterus; vd, vas deferens.

FIGURE 42. Radulae. A–C. Trochomorpha rubens, FMNH 175247. D–F. Videna planorbis, FMNH 160957. G–I. Dendrotrochus eva eva, FMNH 15719. J–L. Orpiella concavum, BMNH no registration number. Scale bars 20 mm (A–L).

FIGURE 44. Trees resulting from the phylogenetic analysis of two molecular datasets. Families and subfamilies are shaded. A. Mitochondrial gene cytochrome c oxidase subunit I [after Hyman et al. (2007)]. B. Nuclear gene 28S rRNA [after Wade et al. (2006)].