Kalidos balstoni (Angas, 1877)

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 : 89

Kalidos balstoni (Angas, 1877)

Figures 23G–H , 29E , 31M–O , 33B , 35A

Material examined. Madagascar: AM C205303 (two specimens dissected, one radula examined), 15km SW of Andringitra Reserve, 55 km SE of Ambalabao , SW Madagascar, dense mountain rainforest, 1997 .

Description. External morphology: Shell 5.9 whorls, orange brown, spire and apex slightly raised. Protoconch sculptured with spiral ribs and irregular radial ribs; teleoconch with radial ribs. Whorl profile flattened above and below an angular periphery. Internal walls of early whorls complete; umbilicus open, very narrow U-shape. Animal ( Figure 35A ) light brownish-yellow with grey spots. Mantle laps absent. Right mantle lobe small, left and median mantle lobes very small. Sole of foot and caudal apparatus as for family; caudal horn small; caudal foss vertical slit in tail.

Mantle cavity: As for family. Mantle with no visible minor blood vessels, pigmentation of black and white spots.

Digestive system: Oesophageal crop absent.

Genital system ( Figures 23G–H , 29E , 33B ): As for family. Carrefour and talon both not embedded in albumen gland. Free oviduct of medium length; capsular gland present; internal longitudinal pilasters absent. Bursa copulatrix inserted on vagina, moderately short, about half spermoviduct length; duct of bursa copulatrix narrow, very short, distinguishable from bursa copulatrix, internally with pustules. Vagina internally with longitudinal pilasters. Penis long; epiphallus enters penis through short verge; penis internally smooth, twelve longitudinal penis pilasters present, penial diverticulum absent. Penial sheath present, very thick, enclosing only penis; muscle connecting top of penial sheath to epiphallus present; penis retractor muscle attached to epiphallus. Epiphallus approximately equal to penis length, internally with longitudinal pilasters. Epiphallic retractor caecum absent. Flagellum present; flagellum and distal part of epiphallus without internal cryptae and externally smooth and slender. Spermatophore soft capsule with firm tail pipe open at one end; tail pipe long, sculptured with longitudinal rows of tiny teeth.

Radula ( Figure 31M–O ): Relatively long and narrow. Central tooth with small ectocones; mesocone lanceolate, similar in length to tooth base. Lateral and marginal tooth fields distinguishable. Lateral teeth with endocone much smaller than central tooth ectocones; ectocone equal in size to those on central tooth; mesocone shorter than tooth base. Marginal teeth with endocones absent; ectocones absent, not subdivided into extra teeth. Radular formula (69.12.1.12.69) × 105 rows.

Remarks. Kalidos is a large genus endemic to Madagascar containing about 60 species ( Schileyko 2002b). No anatomical data are available for the type species, K. ekongensis (Angas, 1877) . The genus was traditionally included in Ariophantinae (Ariophantidae) ( Zilch 1959) but has been more recently regarded as a member of Ereptinae (Ariophantidae) ( Schileyko 2002b). Kalidos balstoni diverged from the base of Helicarionidae in the present phylogenetic analysis ( Figures 1–3 ). In the molecular analysis of Hyman et al. (2007) its position was poorly resolved, but it diverged basally from Helicarionidae when other problematic taxa were removed.

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 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).

FIGURE 29. Reproductive systems. A. Caldwellia angularis, AM C205299. B. Caldwellia imperfecta, AM C205300. C. Erepta odontina, AM C205302. D. Ctenophila caldwelli, AM C205301. E. Kalidos balstoni, AM C205303. Scale bars 1 mm (E), 2 mm (B, D), 5 mm (A, C). Abbreviations: alb, albumen gland; bc, bursa copulatrix; c, caecum; ca, carrefour; ep, epiphallus; fl, flagellum; fov, free oviduct; her, hermaphrodite duct; ov, ovotestis; pr, prostate; prm, penis retractor muscle; pt, penial sheath; ut, uterus; v, vagina; vd, vas deferens.

FIGURE 31. Radulae. A–C. Caldwellia imperfecta, AM C205300. D–F. Caldwellia angularis, AM C205299. G–I. Ctenophila caldwelli, AM C205301. J–L. Erepta odontina, AM C205302. M–O. Kalidos balstoni, AM C205303. P–R. Madagascan species MA1, AM C205304. Scale bars 10 mm (A–C), 20 mm (D–L, P–R), 50 mm (M–O).

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 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).