Landouria intumescens (Martens, 1867) comb. nov.

Figs 42, 53, 66, 72–74; Tables 3–4

Helix intumescens Martens, 1867: 263, pl. 13, fig. 10 (“bei Surabaya und dem benachbarten Grisse … im Tengergebirge bei Wonosari”) (in part?).

Landouria winteriana – van Benthem Jutting 1950: 463 (in part). (not Pfeiffer, 1842)

Diagnosis

Landouria intumescens is characterized by a medium-sized, brownish-corneous, depressed conical, sharply keeled shell and a penis that is divided by a constriction into a long distal and a short proximal section and a flagellum with a broad, crenated distal part and a short and narrow proximal part.

Material examined

Lectotype

INDONESIA • lectotype (here designated); East Java, Surabaya; 7°20′ S, 112°45′ E; E. von Martens leg.; measurements: D = 13.4 mm, H = 8.2 mm; ZMB 5160 a.

Paralectotypes

INDONESIA • 7 spec.; same data as for lectotype; ZMB 5160.

Other material

INDONESIA – East Java • 2 spec. (det. anat.); Tuban, Montong, Guwoterus, teak and agroforestry; 6°57′56″ S, 111°47′56″ E; 122 m a.s.l.; MZB 19208 • 11 spec. (det. anat.); Tuban, Montong, Guwoterus, sumber Krawak, secondary forest; 6°57′50″ S, 111°48′20″ E; 147 m a.s.l.; MZB 19206 • 1 spec.; same data as for preceding; MZB 19207 • 8 spec.; same data as for preceding; ZMH 133438 • 20 spec.; Bojonegoro, Dander, Sumberarum, mouth of Lawa Cave; 7°16′30″ S, 111°51′09″ E; 117 m a.s.l.; MZB 19284 • 10 spec.; same data as for preceding; ZMH 1334110 • 1 spec.; Bojonegoro, Dander, Sumberarum, mouth of Sumur Cave; 7°16′08″ S, 111°51′41″ E; 72 m a.s.l.; MZB 19263 • 4 spec.; same data as for preceding; MZB 19316 • 8 spec.; same data as for preceding; ZMH 133415 .

Description

SHELL (Figs 72–74; Tables 3–4). Depressed conical, with 5–5.75 slightly convex whorls; protoconch almost smooth; teleoconch with fine, irregular wrinkles; apical side with indistinct incised spiral lines, umbilical side with more distinct spiral lines; without scaly processes; tuberculate around umbilicus; brownish-corneous, some with a light brownish band at periphery; body whorl sharply keeled; aperture rounded rhombic; upper insertion of peristome slightly descending; peristome expanded, reflexed and slightly thickened; umbilicus slightly eccentric, wide, comprising 22–34% of shell diameter, hardly obscured by columellar edge.

GENITALIA (Figs 42, 66; Table 4). Atrium short; penis long, distal part cylindrical, broad, tapering proximally, separated from proximal part by a constriction before bend in penis, proximal part slightly dilated at its proximal end, without penis coecum; membranaceous sheath around distal part of penis connected with proximal part of epiphallus by tissue; penial retractor runs from diaphragm to middle part of short, cylindrical epiphallus; flagellum moderately long, broadest near its distal end, slightly crenated, without node, proximal part narrow, almost cylindrical; vas deferens narrow and long; vagina long, distally broad, proximally narrow; oviduct shorter, proximally tapering; peduncle of bursa copulatrix subdivided into a broader, cylindrical distal part and a narrower, longer proximal part, ending in an elongate oval bursa beside albumen gland. Right ommatophoral retractor runs between penis and vagina.

Remarks

Martens (1867: 263, pl. 13, fig. 10) proposed the name H. intumescens for a medium-sized morph from Surabaya, which he figured, but also included a smaller morph from Wonosari in the Tengger Mountains in the nominal taxon. To fix the name for the morph from Surabaya, we designate the specimen figured by Martens (1867: pl. 13, fig. 10; see Fig. 72) as the lectotype.

We examined specimens from Tuban Regency anatomically; they agree in shell characters with the morph from Surabaya. The specific identity of these specimens with L. intumescens must be proven by an anatomical examination of specimens from Surabaya.

Whether the smaller paralectotypes of H. intumescens from Wonosari in the Tengger Mountains (ZMB 230704; Fig. 74) are conspecific with L. intumescens must also be proven by an anatomical examination.

Landouria intumescens cannot be reliably distinguished from L. winteriana by shell characters. However, the shells of L. intumescens usually differ from those of L. winteriana in the larger diameter (11.6– 14.1 mm vs 9.7–12.6 mm in L. winteriana) and in the relatively narrower umbilicus, which comprises 22–34% of the shell diameter in L. intumescens vs 29–51% in L. winteriana . Landouria intumescens also differs from L. winteriana in the penis, which becomes narrow towards its proximal end, whereas it ends proximally in a globular dilatation in L. winteriana (Fig. 15).

Distribution

Landouria intumescens is known only from northern East Java (Fig. 53).