Helix (Chloritis) pseudomiara Bavay & Dautzenberg, 1909 by monotypy.

The anatomically-examined specimen (i.e., on which the genus is based) was in fact Helix condoriana. Under the provisions of Article 70.3 ICZN, we herewith replace the original type species Helix (Chloritis) pseudomiara Bavay & Dautzenberg, 1909 with Helix condoriana Crosse & Fischer, 1863 as the type species of Bellatrachia Schileyko, 2018 to serve the stability of nomenclature.

Bellatrachia condoriana (Crosse & Fischer, 1863).

Shell depressed globular, apex not sunken, hairs or hair scars cover the entire shell. Penis rather long, subcylindrical, its inner surface bears longitudinal pilasters; penial verge absent; penial caecum absent; epiphallus slender, long, convoluted; retractor muscle attached at the penis-epiphallus transition; flagellum thick, with attenuated tip, approximately 2–2.5 times shorter than epiphallus; vagina slender, shorter than penis; stalk of bursa copulatrix long, with thickening at some distance from its origin, shape of bursa unknown (based on Schileyko 2018; see Fig. 4).

See B. condoriana below.

Bellatrachia differs from Trichochloritis in lacking the penial caecum.

condoriana: 1 syntype MNHN-IM-2000-1866, Poulo-Condor, D: 18.3 mm, H: 11.7 mm [this is the syntype figured in the original description]; microtricha: lectotype (selected by Zilch 1966) SMF 8540, Vietnam, Annam, Boloven, coll. O. Möllendorff ex coll. Roebelen, D: 24.8 mm, H: 17.7 mm.

“insula Poulo-Condor” [Con Son Island], Vietnam.

Vietnam, Bang-Kiang, coll. Denis ex Messager, MNHN-IM-2012-27151 (2 shells).

Shell biconvex with a whitish subsutural spiral, narrow umbilicus, and hair scars covering the entire surface.

Shell middle sized, biconvex, moderately thin-walled; last whorl only slightly expanding and descending abruptly towards aperture; colour dirty yellowish with a broad pale subsutural spiral band; whorls 4.5–5, separated by a rather shallow suture; body whorl faintly slightly angled; subsutural furrow shallow but present on the complete last whorl; protoconch consists of 1.25–1.5 whorls, very finely squamous, matte; the pattern of hair scars is dense and covers the complete teleoconch; aperture obliquely rounded, and the peristomal rims are close; peristome strongly expanded and somewhat reflected and reinforced by a white lip; parietal side with very thin, inconspicuous light layer; umbilicus open, of medium size, with blunt peripheral angulation, and partly covered by the columellar reflection.

D = 18.3–24.8 mm; H= 11.7–17.7 mm (n = 4).

The syntype of B. condoriana (Fig. 1) is similar to the specimen identified as Helix (Chloritis) pseudomiara by Schileyko (2017) (Fig. 3), but the shell of the latter is somewhat more depressed. The shell of the lectotype of B. microtricha (Fig. 2) is larger and somewhat more globular that that of B. condoriana. However, both taxa agree quite well in other details such as the relative size of the umbilicus, formation of lip and aperture, and microsculpture of the teleoconch. In contrast, absolute dimensions proved to be insufficient traits for species-level distinction. Therefore, we consider Chloritis microtricha as a synonym of Bellatrachia condoriana. The subtle conchological differences in the shell morphology shown in Figs 1–3 may be part of the overall variation of B. condoriana or may signal a difference at the species level. This question can only be clarified by a revision of a larger number of specimens from the area.

Figures 1–2. 

Bellatrachia condoriana 1 Syntype Helix condoriana Crosse & Fischer, 1863, MNHN-IM-2000-1866, D = 18.3 mm, MNHN 2 lectotype Chloritis microtricha Möllendorff, 1898, SMF 8540, D = 24.8 mm, S. Hof, Senckenberg. All photographs × 2.

Figures 3–4. 

Original specimen “Bellatrachia pseudomiara” sensu Schileyko 3 shell (Schileyko 2018: 170, Fig. 1), D = 23.2 mm, A. Sysoev, photograph × 2 4 Morphology of the genital organs of “Bellatrachia pseudomiara” sensu Schileyko (2018), modified after Schileyko (2018).

Helix breviseta L. Pfeiffer, 1862 by original designation.

Helix breviseta L. Pfeiffer, 1862, Trachia penangensis Stoliczka, 1873.

Shell depressed globular, apex not sunken, hairs or hair scars cover the entire shell. Penis thickened, probably with penial verge (?) and a slender, relatively long penial caecum; epiphallus slender, shorter than penis; retractor muscle attached at the penis-epiphallus transition; flagellum short; vagina slender, shorter than penis; stalk of bursa copulatrix long, with thickened base and oval bursa (based on the drawings of Stoliczka 1873: plate 3, fig. 18 and Collinge 1903: plate 12, fig. 17.).

The anatomy of the genital organs of Helix (Trachia) malayana Möllendorff, 1887 (= Trichochloritis breviseta; see Maassen 2001) was described by Collinge (1903), and that of T. penangensis is known from Stoliczka (1873), here re-drawn and provided in Fig. 8 (penangensis) and Fig. 9 (breviseta). Both species possess a penial caecum, which is here considered as a diagnostic trait for the genus. Without knowing the full anatomy, it is uncertain how many of the hairy Chloritis-like species of continental Asia belong to this group.

breviseta: syntype MNHN-IM-2000-1847, Siam, D: 22.1 mm, H: 12.9 mm; malayana: syntypes (2 shells) NHMUK 1891.3.17.3–4, Perak, leg. Hungerford.

Perak, leg. Hungerford, NHMUK 1891.3.17.3–4 (2 shells of “malayana”); Larut, Malay Peninsula, NHMUK 1897.3.15.7 (1 shell of “malayana”); Malakka, Kelantan, Hochland v. Perak, coll. O. Möllendorff ex coll. H. Rolle ex coll. Waterstraat, SMF 8538/1 (“malayana”).

“Siam” (breviseta); “Perak” [Perak state, Malaysia] (malayana).

Shell depressed, unicoloured, yellowish, with permanent hairs; umbilicus funnel-shaped with a blunt peripheral angulation.

Spire only slightly elevated, shell depressed, shell thin; last whorl bluntly angled, a subsutural furrow is present but insignificant; colour yellowish, spiral band missing; the 4.5 whorls separated by a rather shallow suture; protoconch consists of slightly more than 1.5 whorls, squamous, bears minute wrinkled hair scars; teleoconch completely covered by a moderately dense pattern of hairs; bristles stiffy and durable and stick to the shell (their apical part breaks off, but a dark brown conical bristle cone is left making the surface of the shell quite rough); aperture subrectangular with only slightly oblique columella; peristome reflected and covered by a white lip; parietal region with very slight whitish, blunt lime layer, inconspicuous; columellar reflection small; umbilicus wide and funnel-shaped with a blunt peripheral keel.

Figures 5–7. 

Trichochloritis breviseta 5 syntype Helix breviseta L. Pfeiffer, 1862, MNHN-IM-2000-1847, D = 22.1 mm, MNHN 6 syntype Helix (Trachia) malayana Möllendorff, 1887, NHMUK 1891.3.17.3, D = 22.2 mm, NHMUK 7 SMF 8538 ex coll. Möllendorff, D = 20.6 mm, S. Hof, Senckenberg. All photographs × 2.

Figures 8–9. 

Morphology of the genital organs of Trichochloritis species 8 Trichochloritis penangensis (Stoliczka, 1873) (redrawn from Stoliczka 1873) 9 Trichochloritis breviseta (L. Pfeiffer, 1862) (redrawn from Collinge 1903). Not to scale.

Figures 10–12. 

Shells of Trichochloritis species 10 Trichochloritis breviseta, BOR/MOL 9091, Perak, Ipoh, Gunung Kanthan plot, D = 19.5 mm 11 Trichochloritis penangensis, BOR/MOL 11562, Perak, Ipoh, Gunung Pondok, plot, D = 16.2 mm 12 Trichochloritis (?) pseudomiara, syntype of Helix (Chloritis) pseudomiara Bavay & Dautzenberg, 1909, D = 24.3 mm, MNHN. 10, 11 Junn Kitt Foon (published in Foon et al. 2017), all photographs × 2.

D = 22.9–24.1 mm; H = 12.9–14.7 mm (n = 4).

Malaysia and Thailand

The types should be in the Zoological Survey of India in Kolkata but were not found during a recent search (S.K. Sajan, pers. comm., December 2018). They were likewise not found in the NHM.

“Penang”.

“Chloritis penangensis has a much more globular shell with less expanded whorls compared to Chloritis breviseta which has more expanded (perpendicular to the axis) whorls and thus, “wider” looking shells. These characters appear consistent for each species across Peninsular Malaysia (based on conchological comparisons), although shell size varies within each species.” (Junn Kitt Foon, pers. comm., 01 Dec 2018). To illustrate these differences, we illustrated the shells of both species (Figs 10, 11).

Thailand: Krabi: Phanom Benja National Park, Huai To waterfall and surrounding rain forest, 120 m, 08°14'21"N, 098°54'52"E, 08°14'08"N, 098°55'12"E, leg. Hausdorf, 28.07.2010, ZMH 51997/2.

For a detailed description of the shell refer to Sutcharit & Panha, 2010. Our data on the reproductive anatomy largely matches that of Sutcharit and Panha (2010), with the following two exceptions: the flagellum is relatively long and slender, and the penial verge is not irregularly shaped but conical and deeply grooved with the folds starting from the epiphallus.

syntype MNHN-IM-2000-31774, Nat Son, leg. Messager, D: 24.3 mm, H: 13.3 mm.

Muong-Hum, RBINS/1; Muong-Hum, leg. Messager 1908, RBINS/1; Tonkin, Phong-Tho, RBINS/1 (mixed sample with Trichochloritis sp.); Nat-Son, RBINS/3 (mixed sample with Trichochloritis sp.); Tonkin, leg. Messager, RBINS/22 (some of them are juveniles); Tonkin, Phong-Tho, NHMUK 1909.7.9.57/1 (photographed for the North Vietnamese Land Snail Guide); Tonkin, Muong-Hum, MNHN-IM-2012-27105/2; Vietnam, Cam Duong, MNHN-IM-2012-27106/2 (probably erroneous locality?); Tonkin, Phong-Tho, MNHN-IM-2012-27107/2; Tonkin, Muong-Hum, MNHN-IM-2012-27108/2; Tonkin, Muong-Hum, leg. Messager, MNHN-IM-2012-27109/2; Tonkin, MNHN-IM-2012-27110/1; Haut Tonkin, MNHN-IM-2012-27111/1.

Vietnam, N Vietnam: Nat Son.

A rather large, usually dark species with rounded body whorl, fine radial growth lines and deep hair scars; umbilicus open, only a small part of it is covered by the columellar reflection.

Shell rather large, almost flat, with relatively thick wall; body whorl rounded; last half whorl with or without very shallow subsutural furrow; the 4.75–5.25 whorls are separated by a shallow suture; colour greyish yellowish, or brown to olive green; protoconch consists of 1.5 whorls, finely granulate, with fine radial lines near the suture of the last half whorl; teleoconch finely, irregularly wrinkled, and covered with very deep hair scars, which are visible to the naked eye as well on the body whorl; hairs not permanent, although we did not have access to live collected specimens; aperture ovoid; peristome expanded and slightly reflected, and reinforced by a thickened whitish/light brown lip; parietal region with an inconspicuous layer, which is often darker than the rest of the shell; umbilicus widely open, concave and funnel-shaped, slightly covered by reflected peristome.

D = 21.3 –26.0 mm, H = 11.8–14.4 mm (n = 3).

This species is known only from the northernmost part of Vietnam, along the Chinese border.

This species can easily be identified based on the dark green-coloured shell and the deep, widely spaced hair scars that cover the entire teleoconch.

Helix brevidens Sowerby I, 1841: 25 (Puerto Galero, Philippines).

Shell depressed globular, apex not sunken, hairs or hair scars cover the entire shell, aperture with a basal denticle. Penis very thick-walled, with narrow lumen, internally with very large conic tubercles in main chamber; flagellum and epiphallus absent; vas deferens passes gradually enlarging into penis; retractor muscle inserts at curvature of vas deferens close to its joint with penis; penial sheath thin, surrounds upper two third part of penis; vagina shorter than penis, thick.

The name Dentichloritis refers to the presence of a denticle on the basal peristomal lip and the conchological similarity to Chloritis.

There are seven Trichochloritis species known from the Philippines (Richardson 1985), and four of them have been photographed by Zilch (1966). They differ from D. brevidens in the open umbilicus and the lack of denticle on the basal lip. Therefore, we retain them in Trichochloritis until ethanol-preserved specimens become available.

Philippines, m.c. (Museum Cuming), 3 syntypes NHMUK 20190452 (D of photographed shell = 19.5 mm [Fig. 13]).

Philippines, Puerto Galero (Municipality of Puerto Galera, municipality in the province of Oriental Mindoro).

A middle-sized, yellowish species with a slender reddish peripheral belt, short hairs on the entire shell, nearly closed umbilicus (only visible in oblique view), and a slight thickening (denticle) on the basal part of peristome.

Shell medium sized, depressed globular; body whorl rounded with slight indication of a blunt shoulder; last quarter to half whorl with a very shallow subsutural furrow; the 3.75–4 whorls are separated by a shallow suture; colour yellowish to ochre with a reddish slender belt above shoulder (midpoint of body whorl); protoconch consists of 1.5–1.75 whorls, finely granulate, with fine radial wrinkles; teleoconch covered by short hairs or hair scars, which are visible to the naked eye as well; aperture semilunar; peristome expanded and slightly reflected, and reinforced by a thickened whitish brown lip; a slight swelling (denticle) visible on basal part of peristome, between the midpoint of the basal peristome and the columella; parietal region with an inconspicuous layer, which is matter than the rest of the shell; umbilicus nearly closed by columellar reflection, visible only by oblique view.

Figures 13–14. 

Dentichloritis brevidens 13 syntype Helix brevidens Sowerby I, 1841, NHMUK 20190452, D = 19.5 mm 14 Morphology of the genital organs of Dentichloritis brevidens; modified after Schileyko (2007).

Anatomy: Penis very thick-walled, with narrow lumen, internally with short plicae in basal part and very large conic tubercles in main chamber; flagellum and epiphallus absent; vas deferens rather long, evenly thin down to atrium; approximately one third way up it is attached to penis, and after penis is enlarged and fusiform, then in becomes very thin, thread-like, forming a sharp curvature and passes to penis, gradually enlarging; penial retractor attached to curvature of vas deferens and continues as a fine membrane down to middle part of penis; penial sheath thin, surrounds upper two third part of penis. Vagina shorter than penis, thick; spermatheca without visible division to stalk and reservoir, not attending albumen gland and provided with apical ligament (based on Schileyko 2007: 2113–2114, fig. 2032b, c).

Figures 15–17. 

Chloritis (?) bifoveata 15 shell of dissected specimens of Chloritis bifoveata (Thailand: Krabi: Phanom Benja National Park, ZMH 51997) 16 Situs of its genital organs 17 Penial verge of Chloritis bifoveata. A: verge visible from penis lumen; B: starting to penial verge from epiphallus; C: epiphallus opened until penial verge. Scale bar 1 mm.