Landouria tholiformis, Nurinsiyah & Neiber & Hausdorf, 2019

Nurinsiyah, Ayu Savitri, Neiber, Marco T. & Hausdorf, Bernhard, 2019, Revision of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from Java, European Journal of Taxonomy 526, pp. 1-73 : 37-38

Landouria tholiformis sp. nov.

urn:lsid:zoobank.org:act:C083E997-420A-4CAA-8A67-088C02177B83

Figs 7 , 20 , 34 , 51–52 ; Tables 3–4

Diagnosis

Landouria tholiformis sp. nov. is characterized by a brownish-corneous, sharply keeled shell with scaly processes all over the shell, with a bulge at the proximal end of the penis forming a short coecum, and a crenated flagellum, which tapers abruptly at its proximal end.

Etymology

Landouria tholiformis sp. nov. is named after its dome-shaped shell (Latin: ʻtholusʼ = ʻdomeʼ, ʻformaʼ = ʻshapeʼ; used as an adjective).

Material examined

Holotype

INDONESIA • holotype (det. anat.); Gunung Kidul , Tepus, near Poktunggal Beach, agroforestry; 8°08′42″ S, 110°37′52″ E; 137 m a.s.l.; 9 Aug. 2014; A.S. Nurinsiyah, F.L.H. Irsyad, F.J. Sari and E. Nurlaela leg.; D = 8.7 mm, H = 5.2 mm; MZB 17999.

Paratypes INDONESIA • 8 spec.; Yogyakarta, Gunung Kidul, Tepus, Pego Hill , agroforestry; 8°06′10″ S, 110°37′50″ E; 230 m a.s.l.; MZB 19299 .

Description

SHELL ( Figs 7 , 51 ; Tables 3–4 ). Depressed conical, with a dome-shaped spire, with 6–6.25 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; scaly processes all over shell; tuberculate around umbilicus; brownish-corneous; body whorl sharply keeled; aperture rounded rhombic; upper insertion of peristome descending; peristome expanded, reflexed and thickened; umbilicus slightly eccentric, wide, comprising 32–40% of shell diameter, hardly obscured by columellar edge.

GENITALIA ( Figs 20 , 34 ; Table 4 ). Atrium short; penis long, becoming slightly broader proximally, with a bulge at proximal end forming a short coecum; membranaceous sheath around distal part of penis connected with epiphallus by tissue; penial retractor runs from diaphragm to proximal part of cylindrical epiphallus; flagellum moderately long, broadest in its middle part, crenated, without node, abruptly tapering at its proximal end; vas deferens narrow and long; vagina moderately long; oviduct short, broad, proximally tapering; peduncle of bursa copulatrix subdivided into a broader, almost cylindrical distal part that abruptly passes into a narrower and longer proximal part that ends in an elongate oval bursa beside albumen gland. Right ommatophoral retractor runs between penis and vagina.

Remarks

Landouria tholiformis sp. nov. clusters in the phylogeny ( Fig. 1 ) with L. abdidalem sp. nov. (Fig. 56 ). It differs from that species in the more depressed (D/H 1.77–1.88 vs 1.93–2.14 in L. abdidalem sp. nov.), less sharply keeled shell with scaly processes all over the teleoconch ( Fig. 7 ), in the flagellum, which is slightly longer than the short epiphallus, and in having a bulge at the proximal end of the penis (perhaps this is an individual abnormity).

Landouria tholiformis sp. nov. occurs sympatrically with L. sewuensis sp. nov. (Fig. 58), from which it differs in the less sharply keeled shell with a dome-shaped spire, a more distinct granular microsculpture, with scaly processes all over the shell ( Fig. 7 ) and less distinct spiral lines at the umbilical side, the bulge at the proximal end of the penis that forms a short coecum, and a narrow, crenated flagellum. The flagellum of L. sewuensis sp. nov. is broader than that of L. tholiformis sp. nov. and equipped with two rows of tubercles. There is a globular bulge in the distal part of the vagina in L. sewuensis sp. nov. which is absent in L. tholiformis sp. nov.

Distribution

Landouria tholiformis sp. nov. is endemic to the Gunung Sewu karst region in Gunung Kidul Regency (Fig. 52).

Figures

Figs 6–11. Shell sculpture of species of Landouria from Java (umbilical side of shell in 6; apical side in 7–11). 6. L. conoidea (Leschke, 1914), West Java, Bogor (syntype, ZMH 98144). 7. L. tholiformis sp. nov., Yogyakarta, Gunung Kidul, Tepus, near Poktunggal Beach (holotype, MZB 17999). 8. L. madurensis sp. nov., East Java, Madura Island, Sampang, Nepa Monkey lowland rainforest (paratype, MZB 19209). 9. L. pacitanensis sp. nov., East Java, Pacitan, Srau Beach (paratype, ZMH 148510). 10. L. monticola van Benthem Jutting, 1950, West Java, Gunung Cikuray (paratype, SMF 8880). 11. L. smimensis (Mousson, 1848), East Java, Tosari (ZMA 390881). Scale bars: 1 mm.

Figs 15–24. Genitalia of species of Landouria from Java. 15. L. winteriana (Pfeiffer, 1842), West Java, Padalarang, Gunung Masigit (neotype, ZMA 376193a). 16. L. rotatoria (Pfeiffer, 1842), West Java, Gunung Salak, 1166 m a.s.l. (neotype, MZB 19205). 17. L. epiplatia (Möllendorff, 1897)?, West Java, Bogor, Gunung Gede (ZMH 79946). 18. L. nusakambangensis sp. nov., Central Java, Nusa Kambangan Island, nature monument Gligir (holotype, ZMA 412298a). 19. L. petrukensis sp. nov., Central Java, Karangbolong, way to Petruk Cave (holotype, MZB 19203). 20. L. tholiformis sp. nov., Yogyakarta, Gunung Kidul, Tepus, near Poktunggal Beach (holotype, MZB 17999). 21. L. abdidalem sp. nov., Yogyakarta, Kulonprogo, Sabrang Kidul (holotype, MZB 17664). 22. L. madurensis sp. nov., East Java, Madura Island, Sampang, Nepa Monkey lowland rainforest (paratype, MZB 19209). 23. L. sewuensis sp. nov., Yogyakarta, Gunung Kidul, Tepus, Mbajo Hill (holotype, MZB 19255). 24. L. sukoliloensis sp. nov., Central Java, Kayen, near Watu Payung (holotype, MZB 19219). Scale bars: 1 mm.

Figs 29–47. Flagella of species of Landouria from Java. 29. L. winteriana (Pfeiffer, 1842), West Java, Padalarang, Gunung Masigit (neotype, ZMA 376193a). 30. L. rotatoria (Pfeiffer, 1842), West Java, Gunung Salak, 1166 m a.s.l. (neotype, MZB 19205). 31. L. epiplatia (Möllendorff, 1897)?, West Java, Bogor, Gunung Gede (ZMH 79946). 32. L. nusakambangensis sp. nov., Central Java, Nusa Kambangan Island, nature monument Gligir (holotype, ZMA 412298a). 33. L. petrukensis sp. nov., Central Java, Karangbolong, way to Petruk Cave (holotype, MZB 19203). 34. L. tholiformis sp. nov., Yogyakarta, Gunung Kidul, Tepus, near Poktunggal Beach (holotype, MZB 17999). 35. L. abdidalem sp. nov., Yogyakarta, Kulonprogo, Sabrang Kidul (holotype, MZB 17664). 36. L. madurensis sp. nov., East Java, Madura Island, Sampang, Nepa Monkey lowland rainforest (paratype, MZB 19209). 37. L. sewuensis sp. nov., Yogyakarta, Gunung Kidul, Tepus, Mbajo Hill (holotype, MZB 19255). 38. L. sukoliloensis sp. nov., Central Java, Kayen, near Watu Payung (holotype, MZB 19219). 39. L. tonywhitteni sp. nov., Central Java, Taruman Village, near Sendang Sedayu (holotype, MZB 19214). 40. L. nodifera sp. nov., Central Java, Larangan Village, Lanang Spring (holotype, MZB 19227). 41. L. pacitanensis sp. nov., East Java, Pacitan, near Srau Beach (MZB 19231). 42. L. intumescens (Martens, 1867), East Java, Guwoterus (MZB 19207). 43. L. smimensis (Mousson, 1848), East Java, Tosari (ZMA 390881). 44. L. zonifera sp. nov., Yogyakarta, Kulonprogo, Sibolong (holotype, MZB 17727). 45. L. ciliocincta (Möllendorff, 1897), East Java, Malang, Kondangmerak (MZB 19211). 46. L. pakidulan sp. nov., Central Java, Nusa Kambangan Island, nature monument Gligir (holotype, ZMA 417011a). 47. L. menorehensis sp. nov., Yogyakarta, Kulonprogo, Sibolong (holotype, MZB 19201). Scale bar: 1 mm.

Figs 48–51 (opposite page). Shells of species of Landouria from Java. 48. L. parahyangensis sp. nov., West Java, Karangnunggal, surroundings of Pasir Taraje Cave (holotype, MZB 20429). 49. L. nusakambangensis sp. nov., Central Java, Nusa Kambangan Island, nature monument Gligir (holotype, ZMA 412298a). 50. L. petrukensis sp. nov., Central Java, Karangbolong, way to Petruk Cave (holotype, MZB 19203). 51. L. tholiformis sp. nov., Yogyakarta, Gunung Kidul, Tepus, near Poktunggal Beach (holotype, MZB 17999). Scale bar: 5 mm.

Figs 52–55 (opposite page). Distribution of species of Landouria in Java (10′ grid). 52. L. winteriana (Pfeiffer, 1842), L. naggsi sp. nov., L. nusakambangensis sp. nov., L. petrukensis sp. nov., L. tholiformis sp. nov., L. tonywhitteni sp. nov., L. smimensis (Mousson, 1848). 53. L. conoidea (Leschke, 1914), L. abdidalem sp. nov., L. sewuensis sp. nov., L. sukoliloensis sp. nov., L. pacitanensis sp. nov., L. intumescens (Martens, 1867), L. madurensis sp. nov. 54. L. rotatoria (Pfeiffer, 1842), L. nodifera sp. nov., L. parahyangensis sp. nov., L. schepmani (Möllendorff, 1897), L. zonifera sp. nov., L. ciliocincta (Möllendorff, 1897), L. pakidulan sp. nov. 55. L. epiplatia (Möllendorff, 1897), L. monticola van Benthem Jutting, 1950, L. davini Dharma, 2015, L. menorehensis sp. nov., L. dharmai sp. nov., L. moussoniana (Martens, 1867).

Fig. 1. Bayesian 50% majority rule consensus tree based on mitochondrial 16S rDNA sequences of species of Landouria Godwin-Austen, 1918 with Mandarina suenoae Minato, 1978 and Euhadra peliomphala (Pfeiffer, 1850) as outgroups. All species of Landouria in this tree are from Java except L. omphalostoma Páll-Gergely & Hunyadi, 2013 from China and L. timorensis Köhler et al. in press, L. montana Köhler et al. in press and Landouria sp. nov. (= L. winteriana (non Pfeiffer, 1842) in Köhler et al. (in press)) from Timor. Values at the nodes represent Bayesian posterior probabilities (left) and maximum likelihood bootstrap values (right). Numbers after taxon names correspond to MZB and ZMH catalogue numbers, AM catalogue numbers or GenBank accession numbers (see Table 1).

Figs 56–59 (opposite page). Shells of species of Landouria from Java. 56. L. abdidalem sp. nov., Yogyakarta, Kulonprogo, Sabrang Kidul (holotype, MZB 17664). 57. L. madurensis sp. nov., East Java, Madura Island, Sampang, Nepa Monkey lowland rainforest (holotype MZB 21432). 58. L. sewuensis sp. nov., Yogyakarta, Gunung Kidul, Tepus, Mbajo Hill (holotype, MZB 19255). 59. L. sukoliloensis sp. nov., Central Java, Kayen, near Watu Payung (holotype, MZB 19219). Scale bar: 5 mm.