Pseudopomatias amoenus Möllendorff, 1885

Páll-Gergely, Barna, Fehér, Zoltán, Hunyadi, András & Asami, Takahiro, 2015, Revision of the genus Pseudopomatias and its relatives (Gastropoda: Cyclophoroidea: Pupinidae), Zootaxa 3937 (1), pp. 1-49 : 30-34

Pseudopomatias amoenus Möllendorff, 1885

Figs 5 B–D, 6C, 12A–J.

Pseudopomatias amoenus Möllendorff, 1885 : p. 164.

Pseudopomatias amoenus — Möllendorff 1886: pp 177–178.

Pseudopomatias amoenus var. tumidula Möllendorff, 1886 : p. 178.

Fargesia cornea Heude, 1886 : p. 211.

Pseudopomatias amoenus — Gredler 1887: p. 158.

Pseudopomatias amoenus var. tumidula — Gredler 1887: p. 158.

Fargesia cornea — Heude 1890: p. 130, Plate 36, Fig. 12 (shell), 12a (operculum, radula).

Pseudopomatias amoenus — Kobelt & Möllendorff 1897: p 140. (partim).

Pseudopomatias amoenus var. tumidula — Kobelt & Möllendorff 1897: p. 140.

Pseudopomatias amoenus — Kobelt & Möllendorff 1897: p. 140. (partim).

Pseudopomatias fulvus Möllendorff, 1901 : p. 81.

Pseudopomatias amoenus — Kobelt 1902: p. 273, Fig. 59. (partim).

Pseudopomatias amoenus var. tumidula — Kobelt 1902: p. 273.

Pseudopomatias amoenus — Kobelt 1902, p. 273, Fig. 59. (partim).

Pseudopomatias fulvus — Kobelt 1902: p. 273.

Pseudopomatias fulvus — H. Fischer & Dautzenberg 1904: p. 429.

Pseudopomatias fulvus —Dautzenberg & H. Fischer 1908: p. 206, Plate 6, Figs 11–10 .

Pseudopomatias amoenus — Yen 1939: pp 26–27, Plate. 2, Fig. 26.

Pseudopomatias amoenus var. tumidula and “ var. humidula ” (sic!) (synonym of amoenus )— Yen 1939: p. 27, Plate. 2, Fig. 27 (wrong enumeration, in the book it is Fig. 20).

Fargesia cornea — Johnson 1973: p. 26.

Pseudopomatias fulvus — Thanh 2008, p. 9.

Pseudopomatias fulvus —Ðỗ et al. 2010: p. 15.

Diagnosis. A small to large, very variable species in terms of shell shape, peristome thickness and size. It can be recognized by the absence of plicae in the last whorl, the regular, low ribs on the whole shell, even behind the aperture and the rather irregular aperture shape. The shell shape is usually turriform (sometimes rather spindleshaped); peristome very thin in most cases, but can be thickened in some populations.

Description of the type series of amoenus: Shell light brown to greenish, slender turriform or rarely spindleshaped; apex blunt; 6.75–7.5 (n=3), rather bulging whorls are separated by deep suture; protoconch consist of about 1.5 whorls; the sculpture of the first half whorl is smooth, glossy, than becomes regularly, very finely ribbed; teleoconch is rather regularly, finely ribbed without spiral lines; the ribs very low, especially on the body whorl; they become somewhat irregular behind the aperture; aperture rather rectangular with very smoothly angled columellar-parietal transition and somewhat sharply angled parietal-palatal transition; all specimens has a bit deformed round shape; apertural rim slim and weak, not reflexed; parietal callus weak.

Description of the type series of corneus (synonym of amoenus ): Shell yellow, rather turriform with rather blunt apex; 7–7.25 (n=2) bulging whorls are separated by rather deep suture; protoconch consist of one and a half whorls, first whorl rather smooth, the last half whorl is very finely, regularly ribbed (observed in one, not corroded shell); teleoconch regularly, finely ribbed, but ribs becoming lower and somewhat less regular on the last quarter of whorl; no spiral lines visible; aperture rounded with not angled columellar-parietal transition and smoothly angled parietal-palatal transition; apertural rim slightly (MCZ 167038) or strongly (USNM 472343) thickened; the boundary between the two circles is visible, but not conspicuous; the two circles are about of the same strength; parietal callus normally developed but slightly weaker than the apertural rim.

Description of the holotype of fulvus (synonym of amoenus ): Shell brownish, spindle-shaped with pointed apex; 7.75 (n=1), rather bulging whorls are separated by deep suture; the sculpture of the first half whorl is smooth, glossy; the remaining two whorls of the protoconch are regularly, very finely ribbed; teleoconch regularly, finely ribbed with extremely fine spiral lines which are consist of a series of minute tubercles; ribs well-pronounced, even on the neck region, but they are slightly lower there; aperture round with not angled columellar-parietal transition and slighlty angled parietal-palatal transition; apertural rim well-developed, rather thick but the boundary between the circles does not seem to be visible; callus well-developed.

Measurements (in mm). H: 8.5–10, D: 3.5–4.0 ("typical" amoenus , NHMUK 20130387, n=3); H: 7.6–9.0, D: 3.7–4.1 (corneus, n=3); H: 9.3, D: 4.1 (holotype of fulvus ); H: 8.7, D: 3.4 (shell 13 N of Thất Khê, identical with typical fulvus ); H: 10.9–13.2, D: 4.5–5.4 (NHMUK 1910.1.21.4–6, n=3); H: 9.5–11 mm, D: 3.4–4.3 (Kongshan Cun, n=5).

Differential diagnosis. P. maasseni n. sp. has flatter whorls (especially the last), a shallower suture, more elongated shell and every second ribs around the umbilicus are prominent. For comparisons with P. sophiae n. sp. and P. nitens n. sp., see under those species. All these three species mentioned ( P. maasseni n. sp. P. sophiae n. sp. and P. nit ens n. sp.) co-occur with P. amoenus . See also under P. e o s and Table 6 .

Type material. China: Hupei: Patung; coll. Möllendorff, SMF 40296, (lectotype of amoenus ); same data, SMF 40298 (two paralectotypes of amoenus ); same data, SMF 40297 (five adult and two juvenile paralectotypes of amoenus ); China: Hunan Heng-shan-hsien, coll. Möllendorff, SMF 40299, (lectotype of amoenus tumidula ); same data, SMF 40300 (one paralectotype of amoenus tumidula ); Tchen-K’eou, USNM 472343 (syntype of cornea ); China, Se-chuan, Tchen-K’eou, leg. Farges, MCZ 167038 (2 syntypes of cornea ); Tonkin: Manson-Gebirge (mountains), coll. Möllendorff ex Fruhstorfer, SMF 109826 (holotype of fulvus ).

Museum material. Hupei, Badung, J.J. MacAndrew coll, ex Rolle, NHMUK 20130387/6; China, Hunan, Hungerford coll, NHMUK 1891.3.17.838.9; Hupé, Gredler det., NHMW 22924/2; Südchina, Gerstenbrandt coll, NHMW 74649; China, Hupé, coll. Klemm, NHMW 109202/1; China, Patung, Hupei, coll. Naegele, SMF 342542/ 1 (mixed sample with P. nitens n. sp.: SMF 50008); Tonkin, probably leg. Messager, NHMUK 1907.5.20.86–7; Tonkin, Lao Kay, probably leg. Messager, NHMUK 1910.1.21.7–9; Tonkin, Trinh Thuong, probably leg. Messager, NHMUK 1909.7.2.35; Tonkin, Nat Son, probably leg. Messager, NHMUK 1909.7.2.23; Tonkin, Muong Hum, probably leg. Messager, NHMUK 1910.1.21.4–6; Tonkin, Muong Hum, probably leg. Messager, NHMUK 1909.1.2.33; Tonkin, J.J. MacAndrew coll, NHMUK 20130388/2; Tonkin, Lao Kay, probably leg. Messager, E. R. Sykes coll, Acc. no. 1825, NHMUK 20130389/2; Tonkin, That Khé, probably leg. Messager, J.J. MacAndrew coll, ex Rolle 27/11/09, NHMUK 20130390/2; Tonkin, Muong Hum, probably leg. Messager, J.J. MacAndrew coll, NHMUK 20130391/2; Tonkin, J.J. MacAndrew coll, ex Rolle 27/11/09, NHMUK 20130392/2; Tonkin, Lao Kay, probably leg. Messager, E. R. Sykes coll, Acc. no. 1825, NHMUK 20130393/2; Tonkin, Lao Kay, probably leg. Messager, E. R. Sykes coll, Acc. no. 1825, NHMUK 20130394/2; Tonkin, J.J. MacAndrew coll, NHMUK 20130395/2; Tonkin, Phong Tho, probably leg. Messager, J.J. MacAndrew coll, NHMUK 20130396/1; Tonkin, Phong Tho, probably leg. Messager, E. R. Sykes coll, Acc. no. 1825, NHMUK 20130397/1; Cam Duong, Tonkin, leg. Messager, excoll Dr. W. Blume, Dr. Leo P. Rušnov coll, NHMW 109200 (1 juvenile shell); Tonkin, That-Khé, coll. C. Dosch, ex H. Rolle, SMF 192250/2; Tonkin, That-Khé, coll. Jaeckel, SMF 201186 /1; Tonkin, Muong-Hum, coll. C. Dosch ex H. Rolle ex Messager, SMF 192248/3; Tonkin, coll. Staadt 1969, MNHN 2012-26938/2; Tonkin, Trinh-Thuong, coll. Denis 1945, MNHN 2012-26939/6; Tonkin, Muong-Hum, coll. Denis 1945, MNHN 2012- 26940/2 (mixed sample with P. maasseni n. sp.: MNHN 2012-26941); Tonkin, coll. Denis 1945, MNHN 2012- 26945/14 (mixed sample with P. nitens n. sp.: MNHN 2012-26946); Tonkin, coll. Denis 1945, MNHN 2012- 26947/7; Tonkin, leg. Messager, MNHN 2012-26949/16; Tonkin, Pac-Kha, leg. Messager, MNHN 2012-26950/35 (some of them with strange, strong ribs and reflexed peristome); Tonkin, Nat-Son, leg. Messager, MNHN 2012- 26951/16 (1 adult, 15 juveniles); Tonkin, Muong-Bo, Dinh Lu, leg. Messager, MNHN 2012-26954/39 (including several juveniles); Tonkin, Muong-Hum, leg. Messager, MNHN 2012-26956/1 (mixed sample with P. maasseni n. sp.: MNHN 2012-26956); Tonkin, leg. Messager, MNHN 2012-26958/21; Tonkin, leg. Messager, MNHN 2012- 26961/1 corroded shell; Indochine, leg. Messager, MNHN 2012-26964/1; Indochine, leg. Messager, MNHN 2012- 26967/5 damaged shells (mixed sample with P. sophiae n. sp.: MNHN 2012-26967); Indochine, leg. Messager, MNHN 2012-26968/4 (mixed sample with P. sophiae n. sp.: MNHN 2012-26970 and P. nitens n. sp.: MNHN 2012-26969); Indochine, leg. Messager, MNHN 2012-26983/2 shells and few opercula; Indochine, leg. Messager, MNHN 2012-26984/2; Indochine, leg. Messager, MNHN 2012-26986/23; Tonkin, leg. Messager, MNHN 2012- 26989/4; Tonkin, Muong-Hum, between Muong-Hum and the clouds pass, near the latter one, more than 2000 m a.s.l., leg. Messager, MNHN 2012-26990/11; same data, MNHN 2012-26991/3; same data, MNHN 2012-26992/8 (mixed sample with P. nitens n. sp.: MNHN 2012-27022); same data, MNHN 2012-26993/3; same data, MNHN 2012-26994/6; same data, MNHN 2012-26995/7 (with very strong ribs); same data, MNHN 2012-26996/4; same data, MNHN 2012-26997/11; same data, MNHN 2012-26998/5; same data, MNHN 2012-26999/6; same data, MNHN 2012-27000/8; same data, MNHN 2012-27001/2; same data, MNHN 2012-27002/1; same data, MNHN 2012-27003/45; Tonkin, Muong-Hum, leg. Messager, MNHN 2012-27004/132; Indochine, leg. Messager, MNHN 2012-27005/143 (mixed sample with P. nitens n. sp.: MNHN 2012-27005); Tonkin, leg. Messager, MNHN 2012- 27007/78; Tonkin, leg. Messager, MNHN 2012-27008 (at least 300 juvenile shells, some of them seems to be P. nitens n. sp.); Tonkin, Pakhé, leg. Messager, MNHN 2012-27009/263 (mixed sample with P. sophiae n. sp.: MNHN 2012-27010 and P. ni t e n s n. sp.: MNHN 2012-27011); Tonkin, leg. Messager, MNHN 2012-27013/84; Tonkin, leg. Messager, MNHN 2012-27014/47; Tonkin, leg. Messager, MNHN 2012-27015/263 (mixed sample with P. nitens n. sp.: MNHN 2012-27015); Tonkin, Muong-Hum, leg. Messager, MNHN 2012-27017/38; Tonkin, coll. Denis 1945, MNHN 2012-27018/1 (mixed sample with P. sophiae n. sp.: MNHN 2012-26935 and P. ni t e n s n. sp.: MNHN 2012-26936); Tonkin, Muong-Hum, NHMUK 1909.7.2.30/1 (labelled as "co-type" of messageri); Tonkin, Nat-Son, 1909.7.2.27–28/2 (labelled as "co-type" of latebasis); Tonkin, Muong-Hum, NHMUK 1909.7.2.31/1 (labelled as "co-type" of foveolata).

New material: Vietnam, Ninh Binh, Prov., Cuc Phuong Nat. Park, 2 km road Xom Bong to HQ, 20.21°N, 105.54°E, primary forest, limestone outcrops in thick clayey soil. Handpicked + soil sample, leg. JJ Vermeulen & L. Deharveng, 10.10.1998, NHMUK 1999.1.4.27/1; China, Hubei, Enshi Tujiazu Miaozu Zizhizhou, Enshi Shi, Mufu Cun SW 3 km, Enshi Daxiagu, exit of the stone path, 500 m, leg. Hunyadi, A., 5.11.2010, HA/1; China, Sichuan, Dujiangyan Shi, Taian Zhen, Qingchenghoushan, Cuiyinghu to upper station of Jinli cable station, 1273 m, 30°56.27110'N, 103°28.75198'E, leg. Nakahara, Y., Ohara, K., Okubo, K. & Otani, J. U., 0 9.07.2007., OK/1; China, Sichuan, Dujiangyanshi, Qingchengshan Zhen, below Woyunsi, 1110 m, N30°54.33919', E103°33.40153', leg. Nakahara, Y., Ohara, K., Okubo, K. & Otani, J. U., 10.07.2007., JUO/1, OK/1; Vietnam, Lạng Sơn Prov., Na Sầm 12 km towards Thất Khê, left side of the road 210 m, 22°07.870'N 106°35.038'E, leg. Hunyadi, A., 12.11.2011, HA/1; Vietnam, Lạng Sơn Prov., Đèo Lũng Ph ầy, Thất Khê N 13 km, 475 m, 22°20.363'N, 106°27.098'E, leg. Hunyadi, A. 15.11.2011., HA/2; China, Yunnan, Yiliang Xian, Jiuxiang Fengjingqu, 1818 m, 25°04.27306'N, 103°22.87032'E, leg. Nakahara, Y., Ohara, K., Okubo, K. & Otani, J. U., 10.06.2005., OK/1, PGB/ 1; China, Yunnan, Wenshan Zhuangzu Miaozu Zizhizhou, Guangnan Xian, Shuguang Xiang, Kongshan Cun, 1556 m, 23°42.950'N, 105°04.570'E, leg. Hunyadi, A., 23.03.2011, HA/33, PGB/3; Northern Vietnam, Plateau de Ta Phin, Nam Cay a' Tsin Hō (= Sin Ho), leg. Saurin, MNHN 2012–27019/1; China, Yunnan, Honghe Hanizu Yizu Zizhizhou, Pingbian Miaozu Zizhixian, Pingbian Daweishan Yuanshi Senlin Gongyuan, 2085 m, 22°54.835'N, 103°41.958'E, leg. Hunyadi, A., 27.03.2011., HA/3; Vietnam, Son La Prov., ca. 32 km from Moc Chau to Hanoi (old road), 20°47.351'N, 104°50.063'E, leg. Hemmen, Ch. & H., 0 2.04.2010., HE/1; same data, 0 7.10.2010., HE/1; Vietnam, Lai Chau Prov., ca. 2 km from Patan to Phong To (right off road), 22°28.493'N, 103°12.881'E, leg. Hemmen, Ch. & H., 0 8.10.2011., HE/1; Vietnam, Cao Bang Prov., ca. 34.5 km from Cao Bang to Dong Khe (left off road), 22°27.439'N, 106°24.994'E, leg. Hemmen, Ch. & H., 26.03.2010., HE/1.

Remarks. In our understanding, Pseudopomatias amoenus is a very widely distributed species that has several local morphologically distinct forms. Here we handle Fargesia cornea and Pseudopomatias fulvus as synonyms of P. amoenus , but provide the description of the type series.

Yen (1939) considered P. amoenus tumidula as the synonym of P. amoenus . The lectotype of P. amoenus tumidula is more corpulent than P. amoenus specimens from Badong, but the paralectotype does not differ from them. The location of the type locality of P. amoenus tumidula is questionable because Hêng-shan-hsien (Hengshan Xian) is situated in eastern Hunan (part of Hengyang Shi), whereas Gui-yang-dshou (Guiyang Xian) is in southeastern Hunan (part of Chenzhou Shi). The two localities are about 130 km from each other (Szekeres M., pers. comm.). P. amoenus was recorded from Nanling Guojia Gongyuan (National Park) in Northern Guangdong by Hu et al. (2004). We have not seen the specimens collected there; therefore this information could not be verified.

The specimens collected 13 km north from Thất Khê are identical with the holotype of P. f u l v us. Typical fulvus has thicker peristome than most Vietnamese and Chinese P. amoenus populations. The species was described from the Manson (Mau Son) Mts., eastern Lang Son Province, Vietnam. We collected the species in two localities in northern Lang Son. We were unable to locate the locality published by Dautzenberg and Fischer (1908) (Deux- Ponts= "two bridges").

Ðỗ et al. (2010) reported P. fulvus from the Xuan Son National Park (Phu Tho Province, Vietnam). We received photos of three specimens from Dr. Đỗ Văn Nhượng. Those specimens are similar to the North Vietnamese shells in terms of shell shape and rib density, but have much thicker peristome and the ribbing on the last quarter of whorl is more widely-spaced than on the preceding areas.

Specimens from Jiuxiang Fengjingqu ( Fig. 12 C) are very small, spindle-shaped; ribs blunt, very regular on the whole shell, aperture large, deformed rounded, apertural margin slim. Specimens from Kongshan Cun ( Fig. 12 D) are tall, elongated turriform; ribs sharp, very regular on the whole shell, aperture large, deformed rounded, apertural margin slim. Most shells collected along the Vietnamese-Chinese border ( Figs 12 H–J) are very large, turriform, sometimes with extremely fine spiral structure between the ribs is visible; peristome with angled parietal-columellar transition, apertural margin slim.

In museum collections and newly collected material Pseudopomatias amoenus was frequently in mixed lots with P. maasseni n. sp., P. n i t e ns n. sp. and P. s o ph i a e n. sp. In the Cuc Phuong National Park P. amoenus lives sympatrically with Vargapupa oharai n. sp.

Distribution ( Fig. 11 ): The species inhabits a large geographic area of over 1300 km from Northern Vietnam (Ninh Binh Province) until the Chinese Chongqing and Hubei Provinces.

Figures

FIGURE 5. SEM images of the protoconch (A – B) and operculum (C, E – F: outer side, D: inner side) of Pseudopomatias sp. A. P. nitens n. sp., HNHM 98834; B. P. amoenus Möllendorff 1885, China, Yunnan, Yiliang Xian, Jiuxiang Fengjingqu, 1818 m, 25 ° 04.27306 ' N, 103 ° 22.87032 ' E, leg. Nakahara, Y., Ohara, K., Okubo, K. & Otani, J. U., 10.06. 2005., coll. PGB; C. P. amoenus, Indochine, leg. Messager, MNHN 2012 - 26983; D. P. amoenus, Tonkin, Muong-Hum, between Muong-Hum and the Clouds Pass (Col de Nuages), near the latter one, more than 2000 m a. s. l., leg. Messager, MNHN 2012 - 26997; E – F. P. e o s, Nantou Xian, Renai Xiang, Meifeng-Songgang road, around km 14, 2120 m asl., 24 ° 05.103 ' N 121 ° 10.257 ' E, leg. Hunyadi, 20.10. 2014.

FIGURE 11. Distribution of Pseudopomatias species. Filled circle: P. pe g ue ns i s (Theobald, 1864); open triangle, top up: P. nitens n. sp.; open circle: P. amoenus Möllendorff 1885 s. l.; empty square: P. e o s Pilsbry & Hirase 1905; script " Shan States ": Pseudopomatias shanensis n. sp. The meaning of the numbers: 1: Sandoway, 2: Pegu, 3: locality of P. amoenus near Jiuxiang Fengjingqu, 4: locality of P. amoenus near Kongshan Cun, 5: Northern Vietnamese and Chinese localities of P. amoenus and P. nitens n. sp., 6: Mau Son, That Khé and the new localities of " typical fulvus " (= P. amoenus), 7: Xuan Son National Park; 8: Cuc Phuong National Park, 9: Chengkou (type locality of Fargesia cornea), 10: locality near Mufu Cun (new locality of typical P. amoenus), 11: Badong (type locality of P. amoenus), 12: Changyang (locality of P. nitens n. sp.), 13: Hengshan (possible position of the type locality of P. amoenus var. tumidula), 14: Guiyan Xian (possible position of the type locality of P. amoenus var. tumidula), 15: Nanling Guojia (see Hu et al. 2004).

FIGURE 12. Shells of Pseudopomatias amoenus Möllendorff 1885. A. lectotype of P. amoenus, SMF 40296; B. lectotype of P. amoenus tumidula, SMF 40299; C. China, Yunnan, Jiuxiang Fengjingqu, coll. PGB; D. China, Yunnan, Kongshan Cun, 1556 m, coll. PGB; E. holotype of P. fulvus, SMF 109826; F. Syntype of Fargesia cornea, USNM 472343; G. Syntype of F. cornea, MCZ 167038; H. Tonkin, Muong Hum, probably leg. Messager, NHMUK 1910.1.21.4 – 6; I. Tonkin, Muong-Hum, between Muong-Hum and the clouds pass, near the latter one, more than 2000 m a. s. l., leg. Messager, MNHN 2012 - 26995; J. Tonkin, leg. Messager, MNHN 2012 - 27007. Photos: A, B, E: S. Hof; C, D, G, I, J: B. Páll-Gergely; F: B. Murphy; H: H. Taylor. Scale bars = 5 mm; the smaller one refers to the photos of the whole shell, the larger one refers to the photos showing the lateral and basal side of the shells.