Gudeodiscus (Gudeodiscus) phlyarius (Mabille, 1887)

Pall-Gergely, Barna, Hunyadi, Andras, Ablett, Jonathan, Lương, Hao Văn, Fred Naggs, & Asami, Takahiro, 2015, Systematics of the family Plectopylidae in Vietnam with additional information on Chinese taxa (Gastropoda, Pulmonata, Stylommatophora), ZooKeys 473, pp. 1-118 : 29-37

Taxon classification Animalia Stylommatophora Plectopylidae

Gudeodiscus (Gudeodiscus) phlyarius (Mabille, 1887) Figures 4 A–F, 5 A–C, 9 C–D, 10 C–F, 11 K–X, 12 A–M, 21, 22, 28A, 28C, 31C, 35 J–L

Plectopylis phlyaria Mabille 1887a, Molluscorum Tonkinorum diagnoses: 6. [type locality not specified].

Plectopylis phlyaria . Mabille 1887b, Bulletin de le Société Malacologique de France, 4: 100-101., Plate 2, Figs 1-3.

Plectopylis phlyaria , - Pilsbry 1893, Manual of Conchology..., 2(8): 158, Plate 43, Figs 40-42.

Plectopylis phlyaria , - Gude 1897b, Science Gossip, 4: 139., Figs 61 a–b. [ “Tonkin”].

Plectopylis (Endoplon) phlyaria , - Gude 1899c, Science Gossip, 4: 148.

Plectopylis (Endoplon) phlyaria , - Gude 1899d, Science Gossip, 6: 175.

Plectopylis (Endoplon) phylaria , - Gude 1901c, Journal of Malacology, 8: 113-115., Figs 3 a–f. ["Than Moi"].

Plectopylis (Endoplon) moellendorffi 1901c Gude, Journal of Malacology, 8: 115-116., Figs 4 a–f. [ “Than-Moi”].

Plectopylis gouldingi Gude 1909, syn. n., Proceedings of the Malacological Society of London, 8: 215, 217., Plate 9, Figs 1, 1 a–b. [ “Nat-Son”].

Plectopylis verecunda Gude 1909, syn. n., Proceedings of the Malacological Society of London, 8: 215, Plate 9, Figs 3, 3 a–b. [ “Phony-Tho”].

Plectopylis fallax Gude 1909, syn. n., Proceedings of the Malacological Society of London, 8: 217, Plate 9, Figs 6, 6 a–b. [ “Muong-Bo”].

Plectopylis anterides Gude 1909, syn. n., Proceedings of the Malacological Society of London, 8: 216, Plate 9, Figs 2, 2 a–b. [ “Pac-Kha”].

Gudeodiscus phlyarius phlyarius (and Plectopylis moellendorffi is synonym), - Páll-Gergely & Hunyadi 2013, Archiv für Molluskenkunde, 142 (1): 25-28., Figs 31, 61 a–b, 63-65, 75 (map) 77 a–b, 112-114.

Gudeodiscus fallax , - Páll-Gergely & Hunyadi 2013, Archiv für Molluskenkunde, 142 (1): 8.

Gudeodiscus gouldingi , - Páll-Gergely & Hunyadi 2013, Archiv für Molluskenkunde, 142 (1): 8.

Gudeodiscus verecundus , - Páll-Gergely & Hunyadi 2013, Archiv für Molluskenkunde, 142 (1): 8.

Gudeodiscus phlyarius werneri Páll-Gergely in Páll-Gergely & Hunyadi 2013, syn. n., Archiv für Molluskenkunde 142 (1): 13: Figs 32, 28-29, 34: Figs 76 a–d.

Types examined.

Tonkin, Muong-Bo, leg. Messager, NHMUK 1922.8.29.58 (holotype of fallax , Figure 5C); Tonkin, Nat-Son, leg. Messager, NHMUK 1922.8.29.56 (holotype of gouldingi , Figure 4E); Tonkin, Pac-Kha, NHMUK 1922.8.29.57 (holotype of anterides , Figure 4F); Tonkin, MNHN 24581 (2 syntypes of phlyaria , Figure 4A); Tonkin, Than-Moi, collection Möllendorff ex Fruhstorfer, SMF 150125a (lectotype of moellendorffi , Figure 4B); Tonkin, Than-Moi, collection Möllendorff ex Fruhstorfer, SMF 150125b (paralectotype of moellendorffi ); Tonkin, Phony-Tho, leg. Messager, NHMUK 1922.8.29.55 (holotype of verecunda , Figure 5B).

Museum material examined.

fallax -like shells. Tonkin, région de Lao Kay, coll. Dosch ex Rolle, SMF 172081/4; Tonkin, Muong-Bo, coll. Dosch ex Rolle, SMF 172077/2; Tonkin, Muong-Kong, coll. Dosch ex Rolle, SMF 172080/4; Muong-Hum, leg. Messager, MNHN-IM-2012-2130/1; Pakhé, leg. Messager, MNHN-IM-2012-2132/19; Pakhé, leg. Messager, MNHN-IM-2012-2135/6; Muong-Kong, leg. Messager, MNHN-IM-2012-2138/2; Muong-Kong, leg. Messager, MNHN-IM-2012-2140/3; Muong-Hum, leg. Messager, MNHN-IM-2012-2144/1; Ban-Lao, leg. Messager, MNHN-IM-2012-2146/28; Trinh-Thuong, leg. Messager, MNHN-IM-2012-2147/29; Pakhé, leg. Messager, MNHN-IM-2012-2148/3 ("var. major "); Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2155/6 ("var. major "); Pac-Kha, leg. Messager, MNHN-IM-2012-2208/3 ("var. major "); Ban-Lao, leg. Messager, MNHN-IM-2012-2150/22; Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2157/19; Muong-Kong, leg. Messager, MNHN-IM-2012-2158/10; Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2160/22; Trinh-Thuong, leg. Messager, MNHN-IM-2012-2161/28; Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2168/14; Muong-Kong, leg. Messager, MNHN-IM-2012-2169/10; Muong-Hum, leg. Messager, MNHN-IM-2012-2174/1; Muong-Bo, leg. Messager, MNHN-IM-2012-2178/8; Pac-Kha, leg. Messager, MNHN-IM-2012-2180/4; Tonkin, leg. Messager, MNHN-IM-2012-2182/20; Muong-Hum, leg. Messager, MNHN-IM-2012-2190/7; Long-Ping, leg. Messager, MNHN-IM-2012-2192/11; Long-Ping, leg. Messager, MNHN-IM-2012-2206/16; Pac-Kha, leg. Messager, MNHN-IM-2012-2209/4; Pac-Kha, leg. Messager, MNHN-IM-2012-2210/2; Muong-Kong, leg. Messager, MNHN-IM-2012-2244/1; Pakhé, leg. Messager, MNHN-IM-2012-2245/9; Cao-Bang, leg. Messager, MNHN-IM-2012-2470/2; Na-Ri, leg. Messager, MNHN-IM-2012-2463/1; Col de Nuages, leg. Messager, MNHN-IM-2012-2451/6; Tonkin, leg. Messager, MNHN-IM-2012-2450/15; Nat-Son, leg. Messager, MNHN-IM-2012-2445/1; Tonkin, leg. Messager, MNHN-IM-2012-2442/2; Bac-Kan, leg. Messager, MNHN-IM-2012-2247/1; Nga-Son, leg. Messager, MNHN-IM-2012-2255/1; Environs de Yen Bai, ex coll. labo. de Géologie de la Sorbonne ( entrée 1952), MNHN-IM-2012-2272/1; Pakhé, leg. Messager, MNHN-IM-2012-2340/12; Tonkin, leg. Messager, MNHN-IM-2012-2395/2; Tonkin, leg. Messager, MNHN-IM-2012-2396/2; Muong-Bo, coll. Staadt, 1969, MNHN-IM-2012-2406/4; Tonkin, coll. Letellier, 1949, MNHN-IM-2012-2410/1; Tonkin, coll. Staadt, 1969, MNHN-IM-2012-2412/1; Trinh-Thuong, coll. Staadt, 1969, MNHN-IM-2012-2416/5; Tonkin, coll. Staadt, 1969, MNHN-IM-2012-2420/1; Trinh-Thuong, coll. Lavezzari, 1929, MNHN-IM-2012-2421/10; Tonkin, Pac-Kha, NHMUK 1916.3.16.14/1; Tonkin, Trinh-Thuong, 5/1/09, NHMUK 20130621.1-2/2; Tonkin, Pac-Kha, 14/6/10, NHMUK 20110289/3 (labelled as " anterides "); Tonkin, Pac-Kha, coll. Preston, 3/11/08, NHMUK 20110290/2 (labelled as " moellendorffi "); Tonkin, Muong-Bo, coll. Salisbury ex Beddome, NHMUK 20110291/3 (labelled as " fallax = moellendorffi "); Tonkin, Lao Kay, NHMUK 1920.1.20.15-16/2; Tonkin, Muong-Bo, NHMUK 1909.3.14.18-20/3; Tonkin, Trinh-Thuong, coll. Rosen ex Messager, NHMW 71640/O/9481/1; Tonkin, Haut-Tonkin, Region de Lao-Kay, coll. Rolle, NHMW 92564/2; Tonkin, Muong-Kong, coll. Rušnov ex Rolle ex Messager, NHMW 92565/1; Tonkin, Pac-Kha, NHMW 46226/1; Tonkin, Long-Po (?), coll. Oberwimmer ex Wagner ex Messager, NHMW 92579/1; Tonkin, Muong-Bo, NHMW 46291/2.

gouldingi / anterides -like shells. Pakhé, leg. Messager, MNHN-IM-2012-2133/53; Muong-Kong, leg. Messager, MNHN-IM-2012-2141/14; Na-Ri, leg. Messager, MNHN-IM-2012-2152/8; Nat-Son, leg. Messager, MNHN-IM-2012-2153/118; Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2156/4; Pac-Kha ( Pakhé), leg. Messager, MNHN-IM-2012-2164/44; Bac-Kan, leg. Messager, MNHN-IM-2012-2167/29; Muong-Kong, leg. Messager, MNHN-IM-2012-2170/1; Tonkin, leg. Messager, MNHN-IM-2012-2175/8; Tonkin, leg. Messager, MNHN-IM-2012-2176/10; Muong-Bo, leg. Messager, MNHN-IM-2012-2179/1; Nac-Ri, leg. Messager, MNHN-IM-2012-2187/6; Muong-Hum, leg. Messager, MNHN-IM-2012-2189/1; Long-Ping, leg. Messager, MNHN-IM-2012-2193/1; Bac-Kan, leg. Messager, MNHN-IM-2012-2195/18; Long-Ping, leg. Messager, MNHN-IM-2012-2197/4; Pac-Kha, leg. Messager, MNHN-IM-2012-2200/32; Pac-Kha, leg. Messager, MNHN-IM-2012-2201/15; Tonkin, leg. Messager, MNHN-IM-2012-2203/1; Long-Ping, leg. Messager, MNHN-IM-2012-2207/4; Long-Ping, leg. Messager, MNHN-IM-2012-2213/2; Cho-Ra, leg. Messager, MNHN-IM-2012-2478/1; Bac-Kan, leg. Messager, MNHN-IM-2012-2476/2; Trinh-Thuong, leg. Messager, MNHN-IM-2012-2473/6; Bac-Kan, leg. Messager, MNHN-IM-2012-2465/4; Na-Ri, leg. Messager, MNHN-IM-2012-2464/1; Na-Ri, leg. Messager, MNHN-IM-2012-2462/8; Tonkin, leg. Messager, MNHN-IM-2012-2459/1; Pakhé, leg. Messager, MNHN-IM-2012-2454/8; Col de Nuages, leg. Messager, MNHN-IM-2012-2452/15; Nat-Son, leg. Messager, MNHN-IM-2012-2446/1; Col de Nuages, leg. Messager, MNHN-IM-2012-2214/9; Na-Ri, leg. Messager, MNHN-IM-2012-2220/8; Pakhé, leg. Messager, MNHN-IM-2012-2226/5; Tonkin, leg. Messager, MNHN-IM-2012-2228/1; Muang-Kong, leg. Messager, MNHN-IM-2012-2243/7; Nat-Son, leg. Messager, MNHN-IM-2012-2256/12; Phi-Mi, leg. Messager, MNHN-IM-2012-2334/1; Tonkin, leg. Messager, MNHN-IM-2012-2372/3; Muong-Kong, leg. Messager, MNHN-IM-2012-2429/8; Bac-Kan, leg. Messager, MNHN-IM-2012-2433/16; Bac-Kan, leg. Messager, MNHN-IM-2012-2436/1; Tonkin, leg. Messager, MNHN-IM-2012-2422/8; Pakhé, leg. Messager, MNHN-IM-2012-2389/2; Bac-Kan, leg. Messager, MNHN-IM-2012-2404/1; Tonkin, coll. Levazzari, 1929, MNHN-IM-2012-2405/3; Muong-Bo, coll. Staadt, 1969, MNHN-IM-2012-2407/1; Trinh-Thuong, coll. Levazzari, 1929, MNHN-IM-2012-2409/1; Bac-Kan, leg. Messager, MNHN-IM-2012-2438/1; Tonkin, leg. Messager, MNHN-IM-2012-2439/6; Tonkin, Pac-Kha, coll. Kennard, A.S. ex auct. (Gude), NHMUK 20130620/1; Tonkin, Pac-Kha, coll. Salisbury ex Beddome, NHMUK 20110285/1 (" gouldingi var. minor "); Tonkin, Pac-Kha, coll. Preston, 3/11/08, NHMUK 20110286/2; Tonkin, Pac-Kha, coll. Salisbury ex Beddome, NHMUK 20110287/2 (" anterides "); Tonkin, Pac-Kha, coll. Preston, 3/11/08, NHMUK 20110288/2 (" anterides "); Tonkin, Pac-Kha, 1909.3.17.21-23/3 (" anterides "); Tonkin, Long-Ping NHMUK 1916.3.16.3/1 (" anterides "); Tonkin, Pac-Kha, Tonkin, Pac-Kha, NHMUK 1909.3.17.26-28/3; Tonkin, Pac-Kha, coll. Rosen ex Messager, NHMW 71640/O/9478/2; Tonkin, Bac-Kha, coll. Rušnov ex Rolle ex Messager, NHMW 92566/2; Tonkin, Pac-Kha, NHMW 46225/2; Tonkin, Pac-Kha, coll. Wagner ex Messager, NHMW 71640/O/10290/1; Tonkin, Long-Phai, coll. Wagner ex Messager, NHMW 71640/O/10291/1; Tonkin, Pac-Kha, NHMW 92568/1; Tonkin, Pac-Kha, NHMW 46292/2; Tonkin, Bac-Kan, coll. Wagner ex Messager, NHMW 71640/O/10292/1; Tonkin, Bac-Kan, coll. Oberwimmer, NHMW 71640/O/14029/3; Tonkin, Nat-Son, coll. Rušnov ex Messager, NHMW 103354/1.

“Mixed” gouldingi / anterides / fallax samples. Bac-Kan, leg. Messager, MNHN-IM-2012-2171/20; Trinh-Thuong, leg. Messager, MNHN-IM-2012-2181/44; Pakhé, leg. Messager, MNHN-IM-2012-2185/31; Muong-Bo, leg. Messager, MNHN-IM-2012-2211/3; Col de Nuages, leg. Messager, MNHN-IM-2012-2218/25; Col de Nuages, leg. Messager, MNHN-IM-2012-2222/15; Tonkin, leg. Messager, MNHN-IM-2012-2224/13; Tonkin, Pac-Kha, coll. Dosch ex Rolle ex Messager, SMF 172079/4.

phlyarius -like shells. Tonkin, Than-Moi, coll. Jetschin, SMF 207669/6; Tonkin, Than-Moi, coll. Möllendorff ex Fruhstorfer, SMF 150126/10; Tonkin, Chuot-Ki (?), coll. Jaeckel, S. H., SMF 207676/1; Tonkin, coll. Ehrmann ex Fruhstorfer, SMF 150127/2; Tonkin, Than-Moi, coll. Dosch ex Rolle, SMF 172092/4; Tonkin, Than-Moi, coll. Dosch ex Rolle, SMF 172091/4; Tonkin, Than-Moi, coll. Dosch ex Rolle, SMF 172093/2; Tonkin, Than-Moi, coll. Ehrmann ex Fruhstorfer, H., SMF 150138/1+1jb; Than-Moi, leg. Messager, MNHN-IM-2012-2212/5; Long-Phai, leg. Messager, 1901, MNHN-IM-2012-2232/1; Than-Moi, coll. Staadt, 1969, MNHN-IM-2012-2279/4; Tonkin, coll. Weiss, 1901, MNHN-IM-2012-2281/5; Province de Cao Lang, Lang-Son, Ky Lua, coll. Saurin, MNHN-IM-2012-2288/2; Na-Ri, leg. Messager, MNHN-IM-2012-2474/1; Tonkin, leg. Messager, MNHN-IM-2012-2427/3; Tonkin, leg. Messager, MNHN-IM-2012-2431/1; Tonkin, leg. Messager, MNHN-IM-2012-2391/1; Bac-Kan, coll. Staadt, 1969, MNHN-IM-2012-2392/2; Than-Moi, coll. Staadt, 1969, MNHN-IM-2012-2397/5; Than-Moi, coll. Staadt, 1969, MNHN-IM-2012-2398/1; Lang-Son, coll. Letellier, 1949, MNHN-IM-2012-2401/1; Than-Moi, coll. Staadt, 1969, MNHN-IM-2012-2413/8; Tonkin, coll. Denis, 1946, MNHN-IM-2012-2387/4; Tonkin, Pac-Kha, NHMUK 1916.3.16.13/1; Tonkin, coll. Salisbury ex Beddome, NHMUK 20130599/2; Tonkin, Muong-Bo, 3/11/08, NHMUK 20130600/2; Tonkin, 4/11/01/32, NHMUK 20130601/3; Tonkin, Phu Quac Oai, coll. Biggs, H.E.J., Acc. no. 2258, NHMUK 20130602/4; Tonkin, coll. Trechmann, Acc. no. 2176, NHMUK 20130603/2; Tonkin, Than-Moi, leg. Fruhstorfer, H., NHMUK 1901.12.12.206-208/3; Tonkin, "showing immature armature", coll. Gude, G.K, NHMUK 1916.3.15.3/1; Tonkin, coll. Fruhstorfer, NHMW 40850/2; Tonkin, coll. Rušnov ex Blume, NHMW 92562/2; Tonkin, Than-Moi, NHMW 39292/4; Tonkin, Than-Moi, coll. Klemm, NHMW 79000/K/17483/1; Tonkin, Than-Moi, coll. Rušnov ex Rolle ex Messager, NHMW 92580/2; Tonkin, Than-Moi, coll. Rušnov ex Rolle, NHMW 92581/4; Tonkin, Than-Moi, coll. Rolle, NHMW 71640/O/12301/1; Tonkin, Than-Moi, coll. Edlauer, NHMW 75000/E/38490/3; Tonkin, That-Ké, coll. Oberwimmer, NHMW 71640/O/12300/1; Tonkin, coll. Fruhstorfer, NHMW 40851/1; Tonkin, That-Ke, coll. Oberwimmer, NHMW 92560/2; Tonkin, Bac-Khuon, coll. Rolle, NHMW 50857/1 (mixed sample with giardi ).

verecunda -like shells. Phong-Tho, leg. Messager, MNHN-IM-2012-2177/9; Nat-Son, leg. Messager, MNHN-IM-2012-2447/6; Phong-Tho, leg. Messager, MNHN-IM-2012-2443/4; Phong-Tho, leg. Messager, MNHN-IM-2012-2423/4; Lai-Chau, coll. Morlet, MNHN-IM-2012-2424/1; Son-Ma, coll. Fischer, MNHN-IM-2012-2417/1.

New material examined.

fallax -like shells. 2011/125 Lào Cai Province, 1.5 km N of Bắc Ngầm cross, valley on the left side of the road, 155 m, 22°24.149'N, 104°14.462'E, leg. Hunyadi, A., 02.12.2011., HA/1; Vn11-187 Lào Cai Province, ca. 3 km SW of Nhà Văn Hóa, 22°25.513'N, 104°12.194'E, leg. Hemmen, Ch. & J., 04.10.2011., HE/21 (+2 specimens in ethanol, one of them anatomically examined, Figures 21, 28A).

phlyarius -like shells. Vn10-53 Lạng Sơn Province, right off rd. 1B Long Đống to Bình Gia, 21°53.938'N, 106°25.605'E, leg. Hemmen, Ch. & J., 20.3.2010., PGB/3; Vn10-48 Lạng Sơn Province, ca. 6 km SE Bắc Sơn (rd. Bắc Sơn to Nga Hải, left off rd), 21°52.422'N, 106°21.508'E, leg. Hemmen, Ch. & J., 19.03.2010., PGB/3; Vn09-24 Cao Bằng Province, ca. 1 km N of Mã Phục (right side off rd. 3), ca. 575 m, 22°43.938'N, 106°20.527'E, leg. Hemmen, Ch. & J., 23.03.2009., HE/1, PGB/3; Vn10-49 Lạng Sơn Province, ca. 16 km SE Bắc Sơn (rd. Bắc Sơn to Nga Hải, left off rd), 21°50.019'N, 106°18.405'E, leg. Hemmen, Ch. & J., 19.03.2010., PGB/2+2jb; Vn09-18 Lạng Sơn Province, ca. 27 km S of Thất Khê, right side off rd. #4 (Lạng Sơn-Thất Khê), ca. 300 m, 22°07.484'N, 106°35.427'E, leg. Hemmen, Ch. & J., 13.10.2009., PGB/7; Vn09-19 Lạng Sơn Province, ca. 25 km S of Thất Khê, right side off rd. #4 (Lạng Sơn-Thất Khê), ca. 220 m, 22°06.477'N, 106°35.356'E, leg. Hemmen, Ch. & J., 13.10.2009., PGB/2; Vn10-129 Lạng Sơn Province, ca. 58.5 km from Thái Nguyên to Bắc Sơn (right side off road), 21°51.166'N, 106°13.003'E, leg. Hemmen, Ch. & J., 22.10.2010., PGB/1; Vn10-56 Lạng Sơn Province, ca. 7 km from Đồng Mỏ to Văn Quan (left off rd #279), no GPS data, approximate GPS position: 21.696000°N, 106.547271°E, leg. Hemmen, Ch. & J., 21.3.2010., PGB/5; Vn09-16 Lạng Sơn Province, Tân Mỹ (N of Lạng Sơn), temple south of the entrance of village, ca. 240 m, 21°58.891'N, 106°40.265'E, leg. Hemmen, Ch. & J., 12.10.2009., PGB/3; Vn10-128 Lạng Sơn Province, ca. 69 km from Thái Nguyên to Bắc Sơn (right side off road), 21°54.270'N, 106°15.801'E, leg. Hemmen, Ch. & J., 22.10.2010., HE/8, PGB/9; Vn11-154 Lạng Sơn Province, km 47, 1 road # 1B between Văn Quan and Bắc Sơn, 21°52.785'N, 106°26.262'E, leg. Hemmen, Ch. & J., 01.04.2011., HE/6 (also in ethanol); Vn11-155 Lạng Sơn Province, ca. 55 km from Bình Gia to Lạng Sơn on road 1B (no GPS data), leg. Hemmen, Ch. & J., 01.04.2011., HE/11; Vn11-156 Lạng Sơn Province, ca. 10.6 km from Bình Gia to Lạng Sơn on road 1B, 21°53.639'N, 106°25.895'E, leg. Hemmen, Ch. & J., 01.04.2011., HE/70 (one of them is sinistral!), (anatomically examined, Figures 22, 28C, 35 J–L); Vn11-157 Lạng Sơn Province, ca. km. 50 of road 1B, 10 km to Bình Gia, 21°53.911'N, 106°25.664'E, leg. Hemmen, Ch. & J., 01.04.2011., HE/6 (anatomically examined, see Figure 31C); 2011/65 Lạng Sơn Province, Đồng Mỏ 2.5 km towards Văn Quan, right side of the road, 270 m, 21°40.358'N, 106°34.783'E, leg. Hunyadi, A., 10.11.2011., HA/5; 2011/66 Lạng Sơn Province, Đồng Mỏ 4.5 km towards Văn Quan, left side of the road, 330 m, 21°40.828'N, 106°34.531'E, leg. Hunyadi, A., 10.11.2011., HA/23, PGB/2; 2011/67 Lạng Sơn Province, Đồng Mỏ 6 km towards Văn Quan, left side of the road, 390 m, 21°41.034'N, 106°33.618'E, leg. Hunyadi, A., 10.11.2011., HA/20, PGB/2; 2011/68 Lạng Sơn Province, Đồng Mỏ 7 km towards Văn Quan, Vạn Linh cross., left side of the road, 370 m, 21°41.158'N, 106°33.588'E, leg. Hunyadi, A., 10.11.2011., HA/56, PGB/3; 2011/70 Lạng Sơn Province, Lạng Sơn, NNE side of Núi Vọng Phu, 21°51.183'N, 106°44.950'E, leg. Hunyadi, A., 11.11.2011., HA/3; 2011/72 Lạng Sơn Province, 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/86, PGB/2; 2011/73 Lạng Sơn Province, Na Sầm 10 km towards Thất Khê, left side of the road, 190 m, 22°07.530'N, 106°35.381'E, leg. Hunyadi, A., 12.11.2011., HA/27, PGB/2; 2011/74 Lạng Sơn Province, Na Sầm 5.5 km towards Thất Khê, right side of the road, 165 m, 22°05.466'N, 106°35.425'E, leg. Hunyadi, A., 12.11.2011., HA/10; 2011/75 Lạng Sơn Province, Tân Mỹ, tunnel 200 m towards Na Sầm, 210 m, 21°59.110'N, 106°40.077'E, leg. Hunyadi, A., 12.11.2011., HA/19, PGB/2; 2011/76 Lạng Sơn Province, northern edge of Chi Lăng, pass next to the tourist path (N of Đồng Bành), 75 m, 21°34.945'N, 106°30.567'E, leg. Hunyadi, A., 13.11.2011., HA/1; 2011/78 Lạng Sơn Province, Đồng Mỏ 7 km towards Chi Lăng, right side of the road, leg. Hunyadi, A., 13.11.2011., HA/1; 2011/79 Lạng Sơn Province, Đồng Mỏ 5.2 km towards Chi Lăng, right side of the road, 40 m, 21°37.215'N, 106°32.538'E, leg. Hunyadi, A., 13.11.2011., HA/1; 2012/37 Lạng Sơn Province, Đồng Mỏ 2.7 km towards Chi Lăng, right side of the old road, cave, 70 m, 21°38.286'N, 106°33.391'E, leg. Hunyadi, A., 25.05.2012., HA/10; 2012/38 Lạng Sơn Province, Đồng Mỏ 4-5 km towards Chi Lăng, right side of the old road, 65 m, 21°37.479'N, 106°32.730'E, leg. Hunyadi, A., 25.05.2012., HA/6; Vn11-159 Lạng Sơn Province, at km 74.8 on road 1B, Đồng Đăng to Thái Nguyên (8 km S Bắc Sơn), 21°54.543'N, 106°17.298'E, leg. Hemmen, Ch. & J., 02.04.2011., HE/1; Vn11-158 Lạng Sơn Province, ca. 7.5 km foad 1B from Bình Gia to Bắc Sơn, 21°53.908'N, 106°25.661'E, leg. Hemmen, Ch. & J., 01.04.2011., HE/1; Vn09-06 Ninh Bình Province, Cúc Phương Nat. Park, ca. half way from Park Headquarters to Thousand Year Old Tree, left path, ca 510 m, 20°21.366'N, 105°35.513'E, leg. Hemmen, Ch. & J., 03.10.2009., HE/2; MAA10 Ninh Bình Province, Cúc Phương Nat. Park, path to fairy cave, 20°21'N, 105°54'E (approximate GPS position), leg. Vermeulen, J., coll. Maassen, W.J.M., 10.10.1998., NHMUK 19991444/2 + one juvenile/broken shell (marked with no. 3 on Figure 43); same data, WM/3; Vn10-41 Thái Nguyên Province, Temple Chùa Hang (ca. 1 km S of Chợ Chu), 21°54.070'N, 105°38.856'E, leg. Hemmen, Ch. & J., 16.03.2010., HE/3 (marked with no.2 on Figure 43).

Diagnosis.

The species is very variable in terms of shell characters (spire height, presence/absence of the apertural fold, aperture shape, morphology of the parietal and palatal plicae and lamellae, fine morphology of the periostracum folds) between and within traditionally recognized species which are synonymized here. Therefore, it is impossible to give a general diagnosis.

Measurements

(in mm). D = 19.3-20.2, H = 8.8-9.1 (n=3, " fallax ", MNHN 2012-2155); D = 10.6-11.7, H = 4.5-4.7 (n=4, " gouldingi ", MNHN, IM-2012-2164); D = 13.2-13.4, H = 5.9-6 (n=2, " phlyarius ", Vn10-53); D = 14.7-15.5, H = 7.8-8.5 (n=3, " phlyarius ", Vn09-18); D = 12.4-12.7, H = 5.7-5.8 (n=2, " phlyarius ", MAA10); D = 15.5-17.1, H = 7.7-7.8 (n=2, " phlyarius ", Vn10-56); D = 15.8-16.6, H = 8.8-9 (n=3, verecunda , MNHN 2012-2177). The size range is continuous to from typical anterides / gouldingi to fallax var. major (see Figure 16).

Differential diagnosis.

See under Gudeodiscus anceyi , Gudeodiscus emigrans , Gudeodiscus giardi , Gudeodiscus hemmeni sp. n., Gudeodiscus messageri and Halongella fruhstorferi .

Intrasubspecific diversity. Extremely large. Table 9 summarized the conchological differences between newly collected Vietnamese Gudeodiscus phlyarius samples.

Description of the genitalia.

Typical fallax : Two specimens were anatomically examined. Locality: Lào Cai Province, ca. 3 km SW of Nhà Văn Hóa, 22°25.513'N, 104°12.194'E, leg. Hemmen, Ch. & J., 04.10.2011. (Figures 21, 28A);

Penis rather spindle-shaped, very much thickened in the middle; internally with a fine papillated/reticulated structure (proximal part) which gradually becomes a laterally folded structure with flat calcareous granules between the folds; pockets are ar ranged in a rather straight line; epiphallus much shorter than penis, thickest at the penis-epiphallus transition, slowly becoming slimmer towards the vas deferens; penis and epiphallus connected with weak muscle fibres; penial caecum absent in one of the specimens and very small in the other; retractor muscle thick, short, inserts on the small penial caecum (or on the penis-epiphallus transition of the other specimen); vas deferens very long; the proximal section curves within a translucent, straight tube, most convolutions occurring proximally to the vaginal bulb, before becoming a solid, thick tube (until the sperm-oviduct). Vagina long, centrally with well-developed vaginal bulb; vaginal bulb thick-walled, internally with fine reticulated sculpture; distal part of the vagina internally with low, dense, transversal folds; gametolytic sac and diverticulum long, of equal length, extending in parallel; gametolytic sac spindle-shaped, diverticulum of equal thickness throughout.

typical phlyarius : Two specimens were anatomically examined, both contained a few embryos at an early developmental state. Localities: Lạng Sơn Province, ca. 10.6 km from Bình Gia to Lạng Sơn on road 1B, 21°53.639'N, 106°25.895'E, leg. Hemmen, Ch. & J., 01.04.2011. (Figures 22, 28C); Lạng Sơn Province, ca. km. 50 of road 1B, 10 km to Bình Gia, 21°53.911'N, 106°25.664'E, leg. Hemmen, Ch. & J., 01.04.2011. (Figure 31C).

Penis spindle-shaped with thickened middle section; internally with elongated folds of various thickness; this internal ribbed surface also continues in the small penial caecum; retractor muscle short, inserts on the penial caecum; epiphallus shorter and much slimmer than the penis; distally the penis and proximal part of epiphallus bound with connective tissue; vas deferens very long, proximally simple, slim, curved centrally and covered with a sheath distally simple and thickened. Vagina long with well-developed central vaginal bulb; internally the proximal part of the bulb is almost smooth; this sculpture changes to parallelly folded structure in distal direction (Figure 31C); the distal part of the vagina is strongly folded; gametolytic sac and diverticulum of equal length, both being relatively short.

Radula.

See Table 6 and Figures 35 J–L.

Distribution

(see Figure 43). The populations assigned to Gudeodiscus phlyarius inhabit several regions of northern Vietnam (Lạng Sơn, Cao Bằng, Ninh Bình, and along the border region with the Chinese Yunnan Province) and the Chinese Guangxi. A single shell of typical Plectopylis fallax Gude, 1909 was collected in southern Yunnan, very close to the Vietnamese border (Honghe Hanizu Yizu Zizhizhou, Hekou Yaozu Zizhixian, Laofanzhai Xiang, Sierqi N 1.5 km towards Laofanzhai, 155 m, 22°44.637'N, 103°53.782'E, leg. Hunyadi, A., 19.03.2011., HA/1).

Remarks.

Gudeodiscus phlyarius and taxa of similar appearance are one of the most problematical groups in the Plectopylidae . Gude (1909) described six species ( anterides , cyrtochila , fallax , gouldingi , messageri , verecunda ) from the border region of northern Vietnam with the Chinese Yunnan Province. One species, Plectopylis cyrtochila differs from the rest of the species by the smooth, lenticular shell and week peristome and callus. Therefore, it is discussed separately, under the name Gudeodiscus cyrtochilus . In face of the obvious similarities between the remaining five species, Plectopylis messageri and Plectopylis fallax were only compared with Plectopylis moellendorffi , and Plectopylis verecunda was compared with Plectopylis messageri . The shell characters of Plectopylis anterides and Plectopylis gouldingi were only compared with each other. Shells having transitional characters were explained by hybrid origin. Gude (1909) mentions that a specimen of messageri from Pac-Kha might be a hybrid with moellendorffi , and another specimen from the same locality was believed to be a hybrid of anterides and gouldingi . The shell characters distinguishing Plectopylis messageri and the sympatric species referable to fallax , gouldingi and anterides are stable, therefore Gudeodiscus messageri is handled separately from the rest of the taxa.

In the recent revision of the Chinese members of the family ( Páll-Gergely and Hunyadi 2013), Gudeodiscus phlyarius was reported from several localities in Guangxi. Plectopylis moellendorffi Gude, 1901 was synonymized with Plectopylis phlyarius . Gudeodiscus phlyarius werneri was described from two nearby localities near Duan city. All other Chinese Gudeodiscus phlyarius populations were assigned to the nominotypical subspecies. Gudeodiscus phlyarius phlyarius populations were listed in two separate groups based on their appearance, namely " phlyarius -like, mainly flat, small form" and "larger, strongly-built shell (transition to werneri )".

Here we include the following taxa as synonyms of Gudeodiscus phlyarius : anterides Gude, 1909, fallax Gude, 1909, fallax var. major Gude, 1909, gouldingi Gude, 1909, moellendorffi Gude, 1901, verecundus Gude, 1909, werneri Páll-Gergely, 2013. The last taxon was described on the basis of a keel with a light band around the umbilicus, the dissolved anterior lamella, the posteriorly elongated upper and lower ends of the posterior lamella and the parallel, horizontal palatal plicae. All other formerly recognized species ( anterides , fallax , gouldingi , moellendorffi , verecundus ) have two well-developed lamellae and oblique, usually depressed Z-shaped palatal plica, often with Y-like posterior ends. However, this study revealed that Gudeodiscus phlyarius is a widely distributed, very variable species and at this moment we see no good reason to maintain one of the morphologically distinct forms as a subspecies. Consequently, we synonymize Gudeodiscus phlyarius werneri with Gudeodiscus phlyarius .

According to the original description the anterior lamella of gouldingi is simple whereas that of anterides is "provided with buttresses". The upper parietal plica is in contact with the anterior lamella in gouldingi , but the lamella is shorter and free in anterides . Both the upper and lower plicae are shorter in anterides . The first palatal plica of anterides has a descending ridge; the same plica is straight in gouldingi . Additionally, the palatal plicae of anterides are not united by a vertical ridge and are more widely spaced than in gouldingi (the drawings in the original description show the reverse). All of the differences mentioned by Gude (1909) are unstable even within a single sample (assumed to be single population). For example, six shells were opened from a sample collected in Nat-Son (leg. Messager, MNHN-IM-2012-2153, containing 118 " gouldingi " shells). The length of the lower horizontal plica varies greatly, but extends beyond the anterior lamella in the anterior direction in every cases. One specimen had buttresses on the anterior lamella. Two specimens possessed an anterior lamella and the upper horizontal plica united, whereas in the case of four specimens this plica was free. Even among the few shells examined by Gude, he found that shells exhibited transitional character states between anterides and gouldingi . Therefore, these forms cannot be handled as separate species.

In the original description of Plectopylis fallax , Gude (1909) compared it only with Plectopylis moellendorffi . He did not compare Plectopylis fallax either with Plectopylis anterides , or with Plectopylis gouldingi . Based on the material housed in the NHM and the specimens mentioned in Gude’s (1909) paper, Gude received very few shells from Messager. Examining the type specimens of the above-mentioned taxa revealed that besides the difference in size (typical fallax is larger than anterides and gouldingi ), the only distinguishing feature is the simple and free palatal plicae in fallax and the bifurcated and usually connecting plicae of gouldingi (syn: anterides ). The palatal plicae are very variable even within the same sample (see Figures 11) and certainly cannot be used to separate these taxa. Larger shells usually have separated palatal plicae and smaller shells tend to have joint palatal plicae. In addition, the characteristic “nautiliform” shape of typical fallax shells is also not a reliable distinguishing feature from Plectopylis gouldingi / anterides as this trait is also variable across gouldingi and fallax samples.

Based on shell size, most of Messager’s samples in the MNHN can be assigned to three forms (approximately 11-13 mm: gouldingi , 14-16 mm: fallax , 19-21 mm: fallax var. major ). However, the ranges of shell size overlaps within a few samples (see “mixed” samples under the material) and assigning some of these shells to one of the forms is impossible. The size range from typical gouldingi (11 mm) to fallax var. major (21 mm) shows a clinal variation without interruption (see Figure 16). On the other hand, we found one sample where the shells clearly differ from two separate forms, namely six typical " fallax var. major " (D: 18.9-20 mm) and gouldingi (D: 12.4-13.5) shells. Unfortunately, as in other samples, the collection locality is not exact enough to determine if these specimens were sympatric.

The apertural fold is always present on typical Gudeodiscus phlyarius shells, but can be rudimentary or missing in typical anterides / fallax / gouldingi shells. The edge of the periostracal folds has a pointed structure which seems to occur in a spiralling pattern on the shell of most Vietnamese phlyarius specimens, but these are always missing in fallax and gouldingi specimens (this trait is visible only in fresh shells) (Figures 10 C–F). Typical moellendorffi specimens (synonym of phlyarius ) possess a somewhat elevated spire, whereas typical anterides / fallax / gouldingi shells are almost always entirely flat. The only shell character found to be stable within typical Vietnamese Plectopylis phlyarius shells and Plectopylis anterides / fallax / gouldingi shells, however, is the rounded aperture in the former and the elongated aperture in the latter (Figures 9 C–D). Even this difference is found to be variable in Chinese populations. The populations listed as "transitions to werneri " in Páll-Gergely and Hunyadi (2013) have rather elongated aperture, similar to that of typical Vietnamese fallax shells, but have elevated spire and overall similar shell shape to typical Vietnamese phlyarius . Therefore, we refer to anterides , gouldingi and fallax as synonyms of Gudeodiscus phlyarius .

The genital structure of typical fallax and typical phlyarius differ considerably. Namely, the former lacks the penial caecum or has only a very small one, and has a reticulated inner surface of the penis, whereas the latter has a short penial caecum and its penis has parallel folds on the inner wall. The size of the penial caecum however, may not have a strong taxonomic value because it was found to vary largely within species (e.g. Gudeodiscus multispira , see Páll-Gergely and Asami 2014). The sculpture of the wall of the proximal portion of the penis may have a seasonal variability (see under Gudeodiscus villedaryi and in Discussion).

A sample (MNHN 2012-2177) labelled verecunda , which contained 9 shells from the type locality (Phony-Tho) supports the synonymy of the taxon in relation to gouldingi and fallax , and therefore to Gudeodiscus phlyarius . Seven of the shells were typical verecundus with an elevated spire, a strong apertural fold connected to the callus, and an anterior lamella fused to the lower plica; the plica does not extending beyond the lamella anteriorly (confirmed in 3 shells). The two other shells however, have somewhat lower spires, the apertural fold is not connected to the callus and the lower plica is free from the anterior lamella and extended beyond it anteriorly (one of the two shells was opened). These two shells can be interpreted as transitional forms between verecundus and fallax in terms of spire height, apertural fold and parietal plicae/lamellae morphology. Since transitional forms were found between typical verecunda and fallax shells, Plectopylis verecunda can be interpreted as a local form of fallax having elevated spire and fused anterior lamella and lower plica. Therefore, we synonymise Plectopylis verecunda with Gudeodiscus phlyarius .

There are two Vietnamese “forms” of Gudeodiscus phlyarius which differ from all other typical Vietnamese phlyarius shells. One of the morphologically distinct forms inhabits Ninh Bình Province, where we have knowledge of two populations (number 3 on Figure 43). These shells are smaller and comparatively flatter than the usual phlyarius , and have a characteristic “nautiliform” shape, wider umbilicus, with the last whorl leaving the larger part of the penultimate whorl visible. No differences in the lamellae were recognized. The other form is known from one locality in north-western Thái Nguyên Province (number 2 on Figure 43). This has an elevated spire and narrow umbilicus. Only three specimens are known, and two of them were opened. One of the opened specimens had three very weak parietal lamellae (possibly an abnormal character state, similar to that of the holotype of Plectopylis infralevis ), and the second has the anterior lamella and the lower plica fused; the plica did not extends beyond the anterior lamella in the anterior direction.

Two Chinese populations (near Baxianyan, number 1 on Figure 43) have an oblique anterior lamella and an aperture more reflected downwards.