Landouria winteriana (Pfeiffer, 1842)

Figs 2–3, 15, 29, 52; Tables 3–4

Helix winteriana Pfeiffer, 1842: 41 (“ Java ”).

? Plectotropis kraepelini Leschke, 1914: 211, figs 5–7 (“Tjibodas”). Syn. nov.

Helix winteriana – Philippi 1843 in 1842–1845: 23, pl. II.1, fig. 7. — Pfeiffer 1851 in 1849–1853: pl. 95, figs 1–2; 1852 in 1849–1853: 162. — Martens 1867: 264 (in part).

Plectotropis sumatrana var. moussoniana – Leschke 1914: 211 . (not Martens, 1867)

Landouria winteriana – van Benthem Jutting 1950: 463 (in part).

Landouria rotatoria – van Benthem Jutting 1950: 461 (in part). — Smith & Djajasasmita 1988: 394, pl. 2 fig. 10. (not Pfeiffer, 1842)

Landouria cf. rotatoria – Köhler et al. in press: 6, figs 3A, 5 (in part). (not Pfeiffer, 1842)

Diagnosis

Landouria winteriana is characterized by a brownish-corneous, depressed conical, keeled shell with a wide umbilicus, a penis with a large globular bulge at the proximal end and a flagellum consisting of a crenated distal part and a short tapering proximal part.

Material examined

Type material

INDONESIA – West Java • 1 syntype of Plectotropis kraepelini; Cibodas; 6°45′ S, 107°01′ E; ZMH 98416 • neotype of Helix winteriana (designated here) (det. anat.) (voucher for L. rotatoria in van Benthem Jutting 1950); Padalarang, Gunung Masigit; 6°50′ S, 107°26′ E; 500 m a.s.l.; Jan. 1938; W.Ch. Voute leg.; measurements: D = 12.6 mm, H = 6.9 mm; ZMA 376193 a .

Other material

INDONESIA – West Java • 1 spec. (det. anat.); Bogor, Cisarua, Halimun Salak National Park, 1166 m a.s.l.; 6°39′03″ S, 106°27′15″ E; MZB 19235 • 1 spec. (voucher for van Benthem Jutting 1950); Bogor, Gunung Salak; 1200 m a.s.l.; ZMA407749 • 4 spec.; Bogor, Cigudeg, Sipahang Cave (in Gudawang cave complex), agroforestry; 6°27′00″S, 106°31′00″ E; 91 m a.s.l.; MZB 17602 • 7 spec.; Bogor, Ciampea, limestone hill Gunung Cibodas, secondary forest; 6°33′05″ S, 106°41′09″ E; 296 m a.s.l.; MZB 19238 • 98 spec.; same data as for preceding; MZB 19239 • 11 spec.; same data as for preceding; MZB 19240 • 6 spec.; same data as for preceding; ZMH 133425 • 30 spec.; same data as for preceding; ZMH 133426 • 4 spec.; same data as for preceding; ZMH 133427 • 3 spec.; Bogor, Ciampea, limestone hill Gunung Cibodas, inside Si Panjang Cave; 6°33′09″ S, 106°41′13″ E; 357 m a.s.l.; MZB 19279 • 3 spec.; Bogor, Ciampea, limestone hill Gunung Cibodas, Si Ase (Si Gajah) cave mouth; 6°33′08″ S, 106°41′14″ E; 309 m a.s.l.; MZB 19249 • 1 spec. (voucher for P. sumatrana var. moussoniana in Leschke 1914); Ciampea; 6°33′ S, 106°42′ E; ZMH 20426 • 3 spec. (det. anat.); Bogor, Pabaton 25; 6°35′22″ S, 106°47′33″ E; ZMA 369719 • 2 spec. (vouchers for P. sumatrana var. moussoniana in Leschke 1914); Bogor; 6°36′ S, 106°48′ E; ZMH 20425 • 1 spec.; Bogor; 6°36′ S, 106°48′ E; 230 m a.s.l.; ZMA 407751 •> 10 spec. (vouchers for van Benthem Jutting 1950); Bogor; 6°36′ S, 106°48′ E; 250 m a.s.l.; ZMA 407628 • 1 spec. (voucher for van Benthem Jutting 1950); Bogor, botanical garden; 6°36′ S, 106°48′ E; 250 m a.s.l.; ZMA 407750 • 2 spec.; Sukabumi, Jampangtengah, Ciareuy, bamboo plantation; 7°01′49″ S, 106°48′26″ E; 573 m a.s.l.; MZB 19248 • 1 spec. (voucher for van Benthem Jutting 1950); Lake of Cigombong, forest near lake; 6°45′ S, 106°49′ E; ZMA 407800 • 2 spec.; Sukabumi, Benda, Bodogol Nature Reserve at Gunung Pangrango, 800 m a.s.l.; 6°46′40″ S, 106°51′26″ E; ZMH 140032 • 2 spec. (vouchers for van Benthem Jutting 1950); Bogor, Gunung Pancar, 800 m a.s.l.; 6°35′ S, 106°54′ E; ZMA 407790 • 1 spec. (det. anat.); Bogor, saddle between Gunung Pancar and Gunung Paniisan; 6°35′ S, 106°54′ E; 500 m a.s.l.; ZMA 370986 • 5 spec.; Sukabumi, Nyalindung, near Bibijilan waterfall, secondary forest; 7°01′27″ S, 106°54′21″ E; 696 m a s.l.; MZB 19247 • 7 spec.; Sukabumi, Nyalindung, Buniayu cave complex, Cipicung Cave, secondary forest; 7°02′10″ S, 106°54′21″ E; 755 m a.s.l.; MZB 19244 • 7 spec.; Sukabumi, Nyalindung, Buniayu, secondary forest; 7°02′12″ S, 106°54′26″ E; 795 m a.s.l.; MZB 19246 • 3 spec.; same data as for preceding; ZMH 133422 • 1 spec.; Sukabumi, Nyalindung, Buniayu, Altingia (Rasamala) Forest; 7°02′11″ S, 106°54′35″ E; 805 m a.s.l.; MZB 19245 • 1 spec. (det. anat.); Bogor, Megamendung; 6°38′35″ S, 106°55′23″ E; MZB 3618 • 2 spec. (vouchers for van Benthem Jutting 1950); Bogor, S of Cisarua; 6°41′ S, 106°56′ E; 800 m a.s.l.; ZMA 407712 • 2 spec. (vouchers for van Benthem Jutting 1950); same data as for preceding; ZMA 407719 • 1 spec. (voucher for P. sumatrana var. moussoniana in Leschke 1914); Cibodas; 6°45′ S, 107°01′ E; ZMH 20424 • 1 spec.; Padalarang, Rajamandala; 6°50′ S, 107°21′ E; ZMA 413757 • 1 spec. (voucher for van Benthem Jutting 1950); Jampang, limestone rock near Sukanegara; 7°01′ S, 107°22′ E; 1000 m a.s.l.; ZMA 407746 • 4 spec. (vouchers for van Benthem Jutting 1950); Garut; 7°01′ S, 107°25′ E; RMNH 309721 • 6 spec. (det. anat.); Padalarang, Gunung Masigit; 6°50′ S, 107°26′ E; 500 m a.s.l.; ZMA 376193 • many spec. (vouchers for van Benthem Jutting 1950); Padalarang, Gunung Pawon; 6°49′ S, 107°26′ E; 700 m a.s.l.; ZMA 407631 • 27 spec.; Padalarang, Gunung Pawon, shrubs and palm; 6°49′29″ S, 107°26′12″ E; 626 m a.s.l.; MZB 19237 • 28 spec.; Padalarang, Gunung Pawon, shrubs; 6°49′30″ S, 107°26′13″ E; 689 m a.s.l.; MZB 19243 • 15 spec.; same data as for preceding; ZMH 133422 • 1 spec.; Padalarang, Gunung Pawon, agroforestry; 6°49′31″ S, 107°26′16″ E; 697 m a.s.l.; MZB 19236 • 11 spec.; same data as for preceding; MZB 19254 • 4 spec.; Padalarang, bamboo; 6°49′36″ S, 107°27′09″ E; 784 m a.s.l.; MZB 19251 • 3 spec.; Padalarang, Gunung Hawu, agroforestry; 6°50′04″ S, 107°27′31″ E; 835 m a.s.l.; MZB 19291 • 36 spec.; Padalarang, Gunung Hawu, guava plantation; 6°50′07″ S, 107°27′35″ E; 787 m a.s.l.; MZB 19242 • 20 spec.; same data as for preceding; ZMH 133423 • 15 spec.; Padalarang, Gunung Hawu, agroforestry; 6°50′03″ S, 107°27′46″ E; 775 m a.s.l.; MZB 19241 • 8 spec.; same data as for preceding; ZMH 133424 • 4 spec. (vouchers for van Benthem Jutting 1950); Cibuluh near Cisurupan, slope of Papandayan; 7°21′ S, 107°47′ E; RMNH 309739 • 4 spec.; Cibuluh near Cikajang, slope of Papandayan; 7°21′ S, 107°48′ E; RMNH 309762 • 7 spec.; Tasikmalaya, Karangnunggal, surroundings of Kiara Jenggot Cave, agroforestry; 7°39′37″ S, 108°06′55″ E; 234 m a.s.l.; MZB 19250 • 3 spec.; Tasikmalaya, Setiawaras, near Sambawa Cave, agroforestry; 7°35′02″ S, 108°10′58″ E; 361 m a.s.l.; MZB 17789 .

Description

SHELL (Figs 2–3; Tables 3–4). Depressed conical, with 5–5.75 hardly convex whorls; protoconch almost smooth; teleoconch with fine, irregular wrinkles; apical side with indistinct incised spiral lines, umbilical side with more distinct spiral lines; scaly processes sparse, mainly at periphery; tuberculate around umbilicus; brownish-corneous; body whorl keeled; aperture rounded rhombic; upper insertion of peristome descending; peristome expanded, reflexed and slightly thickened; umbilicus slightly eccentric, wide, comprising 29–51% of shell diameter, hardly obscured by columellar edge.

GENITALIA (Figs 15, 29; Table 4). Atrium short; penis long, distal part cylindrical, proximal part globularly dilated; 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, distal part broader, apically with some indentions, apical part tapering towards its apical end; without node; vas deferens narrow and long; vagina long; oviduct short; peduncle of bursa copulatrix subdivided into a broader, rapidly tapering 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

Landouria winteriana and L. rotatoria are the oldest names for species of Landouria from the Indo-Australian Archipelago. Pfeiffer (1842) described both from Java without more exact locality data. Later, these names have been used for many different species from Java and other islands of the Indo-Australian Archipelago. Thus, it is important to clarify their identity. The original description of H. winteriana was based on material in Pfeiffer’s own collection (Pfeiffer 1852 in 1849–1853). Pfeiffer’s collection was acquired by H. Dohrn and his collection in the Stettin Museum was destroyed during the Second World War (Zilch 1967; Dance 1986). We could not locate type specimens of H. winteriana in other museums (e.g., the collection of von dem Busch in the Geosciences Collection of the University of Bremen, SMF, ZMH). Thus, a neotype has to be designated to clarify the taxonomic status and the type locality of H. winteriana .

We designate a specimen from Gunung Masigit near Padalarang in West Java (ZMA 376193a; Figs 2, 21), which agrees with the original description of Pfeiffer (1842) and the figures of Pfeiffer (1851 in 1849–1853: pl. 95, figs 1–2) in the depressed conical shell with an angular periphery and a wide umbilicus, as the neotype of Helix winteriana Pfeiffer, 1842 . This specimen represents a species that is widespread in western Java, which was more intensively studied than other parts of Java in former times (and still today), so it is likely that the neotype actually belongs to the same species as the specimens described by Pfeiffer (1842).

Köhler et al. (in press) recently redescribed L. winteriana under the name L. cf. rotatoria (not Pfeiffer, 1842). The species from Timor named L. cf. winteriana by Köhler et al. (in press) is not closely related to L. winteriana . Although it is more closely related to species of Landouria from Java than to L. timorensis Köhler et al. in press or L. montana Köhler et al. in press from Timor (see Fig. 1; Köhler et al. in press: fig. 2), it is not identical with any of the species described here from Java, but probably represents a new species.

The only available syntype of P. kraepelini (Fig. 3) differs from usual L. winteriana in the smaller shell diameter at three whorls (4.0 mm vs 4.4–5.2 mm in other L. winteriana) and the slightly more pronounced keel. We suppose that P. kraepelini represents a small variety of L. winteriana . However, the specific identity of P. kraepelini and L. winteriana should be checked by an anatomical examination of specimens resembling the syntype of P. kraepelini from Cibodas, its type locality.

Landouria winteriana differs from L. sukoliloensis sp. nov. in the angular body whorl, the not convoluted distal part of the penis, the short epiphallus and the flagellum with a broader distal part with external indentions. This species occurs sympatrically with L. petrukensis sp. nov., from which it differs in the larger size and the angular, less sharply keeled periphery.

Distribution

Anatomically determined samples of L. winteriana are known from Bogor Regency and Padalarang in the mountain region in West Java (Fig. 52). The shells from Tasikmalaya Regency resemble the anatomically determined samples from Bogor Regency and Padalarang. However, their specific identity has to be checked anatomically. In addition, we have anatomically determined a sample from Rakata Island (3 spec. (vouchers for Smith & Djajasasmita 1988), Krakatau Islands, 6°08′46″ S, 105°26′23″ E, MZB 9354).

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Landouria winteriana has also been recorded from Thailand to the Philippines and the Maluku Islands (Martens 1867; Rensch 1932; van Benthem Jutting 1941, 1950; Vermeulen & Whitten 1998; Boonngam et al. 2008). Besides on Java, L. winteriana might occur on Sumatra, but records from other islands refer to other species.