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Clausiliidae of Vietnam (Gastropoda, Stylommatophora)
Hartmut Nordsieck
General remarks
The clausiliid fauna of Vietnam is as rich as that of southern China, but in contrast to that fauna it is relatively well-known. This is mainly due to the activities of French collectors during the first decades of the French colonial government in Indochina (Mabille 1887a, b, 1889, Fischer 1898, Bavay & Dautzenberg 1899 a, b, 1900, 1903, 1909, 1912, 1915, Dautzenberg & Fischer 1905a, b, 1908). These activities were focussed on the mountainous border regions of Tonkin towards China, because several collectors were members of the French army (e. g. Colonel Messager, who collected most clausiliid species of Tonkin). In that time, some of these regions have also been visited by German collectors (Moellendorff 1901). Southern regions of Tonkin have been explored later (until the First World War), but only a few, while the rest of the country (main part of Annam, Cochinchina) remained unexplored. This discrepancy in knowledge continued until the recent decades during which the exploration of Vietnamese clausiliid fauna has been resumed after a long period of standstill (Loosjes & Loosjes-van Bemmel 1949, 1973, Szekeres 1969, 1970, Nordsieck 2002a, b, 2003, 2007, 2010, Maassen & Gittenberger 2007).
For the genus systematics of Vietnamese clausiliids see Ehrmann (1927: descriptions) and Nordsieck (2002a, b, 2007). The species systematics is currently under revision, as can be seen by comparing the following list with former ones (Nordsieck 2002a, b, 2007).
The species which have been recently collected in Vietnam by C. & J. Hemmen, K. Ookubo and S. Panha are marked by an asterisk.
Systematic list
The species (with synonyms, indicated as = ) are listed in alphabetical order; assumed subspecies are added (indicated as +). The species are accompanied by information on their known geographical range (provinces).
Phaedusinae A. J. Wagner 1922
abstrusa (Szekeres 1970)*, Oospira (O.): Hoa Binh, Ninh Binh, Thanh Hoa.
aciculata (Bavay & Dautzenberg 1909), O. (Lindholmiella): Lai Chau.
acrostoma (Bavay & Dautzenberg 1909), Synprosphyma (S.): Lao Cai, Lai Chau.
antibouddah H. Nordsieck 2003*, Oospira (Atractophaedusa): Quang Ninh.
auregani (Bavay & Dautzenberg 1903)*, Liparophaedusa: Lang Son, Bac Kan, Thai Nguyen.
auricoma (Bavay & Dautzenberg 1899), Synprosphyma (S.): Bac Kan.
babeensis (Bavay & Dautzenberg 1899)*, Synprosphyma (S.) (+ crassitesta H. Nordsieck 2011*): Bac Kan, Thai Nguyen.
bavayi H. Nordsieck 2002, Hemiphaedusa (Selenophaedusa): Lao Cai, Lang Son, Bac Kan.
billeti (Fischer 1898)*, Hemiphaedusa (Selenophaedusa) (= falcifera Bavay & Dautzenberg 1899, = callistomella Bavay & Dautzenberg 1900): Ha Giang, Cao Bang, Lang Son, Bac Kan.
bouddah (Bavay & Dautzenberg 1912), Oospira (O.): Quang Ninh.
callistoma (Bavay & Dautzenberg 1899)*, Hemiphaedusa (Selenophaedusa):Cao Bang, Lang Son, Bac Kan.
cervicalis (Bavay & Dautzenberg 1909), Synprosphyma (S.): Lao Cai.
chiemhoaensis (Sykes 1902), Hemiphaedusa (Selenophaedusa) (= lavillei Dautzenberg & Fischer 1905): Ha Giang, Tuyen Quang.
cochinchinensis (L. Pfeiffer 1841), Phaedusa (P.): Da Nang.
coudeini (Bavay & Dautzenberg 1899), Oospira (Formosana): Lang Son.
cuongi Maassen & Gittenberger 2007, Oospira (O.): Thanh Hoa.
dextrogyra (Bavay & Dautzenberg 1909), Oospira (Formosana): Lao Cai.
dichroa (Bavay & Dautzenberg 1899)*, Phaedusa (P.) (+ duporti Bavay & Dautzenberg 1915*): Lang Son, Bac Kan, Ninh Binh, Hoa Binh.
diplochilus (Moellendorff 1901), Hemiphaedusa (Selenophaedusa): Lang Son.
duella (Mabille 1887)*, Dautzenbergiella (D.) (+ apiostoma Moellendorff 1901 = lyteostoma Moellendorff 1901): Lang Son.
duci Maassen & Gittenberger 2007, Oospira (O.) (+ khanhi H. Nordsieck 2011*+ n. subsp. Thanh Hoa*): Son La, Thanh Hoa.
eregia (Szekeres 1969)*, Oospira (O.): Nghe An, Ninh Binh.
fistulata (Bavay & Dautzenberg 1909)*, Synprosphyma (S.): Lao Cai, Ha Giang.
freyi (Bavay & Dautzenberg 1899)*, Liparophaedusa: Bac Kan, Thai Nguyen.
gastrodes (Moellendorff 1901), Synprosphyma (S.): Lang Son.
gastrum (Moellendorff 1901), Synprosphyma (S.) (= ventriculus Moellendorff 1901): Lang Son.
grangeri (Bavay & Dautzenberg 1899)*, Oospira (Siphonophaedusa): Cao Bang, Lang Son.
inanis (Bavay & Dautzenberg 1909), Phaedusa (P.): Lao Cai.
kebavica (Moellendorff 1901)*, Oospira (Atractophaedusa): Quang Ninh.
lemyrei (Bavay & Dautzenberg 1899)*, Hemiphaedusa (Dendrophaedusa): Lang Son, Bac Kan, Thai Nguyen.
leptotesta H. Nordsieck 2011*, Liparophaedusa: Tuyen Quang.
lypra (Mabille 1887)*, Phaedusa (P.) (= houssayi Fischer 1898, = backanensis Bavay & Dautzenberg 1899, = hamonvillei Bavay & Dautzenberg 1899, + eupleura Bavay & Dautzenberg 1899*, + n. subsp. Son La*, + pereupleura H. Nordsieck 2011*, + subgranulosa H. Nordsieck 2003): Cao Bang, Lang Son, Bac Kan, Thai Nguyen, Vinh Phuc, Ninh Binh, Hoa Binh, Dien Bien, Son La, Thanh Hoa, Nghe An.
mairei (Bavay & Dautzenberg 1909)*, Oospira (O.): Lao Cai, Lai Chau.
mansuyi (Dautzenberg & Fischer 1908), Dautzenbergiella (Mansuyiella): Cao Bang.
margaritifera (Bavay & Dautzenberg 1909), Hemiphaedusa (Margaritiphaedusa): Lao Cai.
micropaviei H. Nordsieck 2011*, Phaedusa (P.): Son La.
mirabilis H. Nordsieck 2003, Thaumatoptyx?: Lao Cai.
miranda (Loosjes & Loosjes-van Bemmel 1973)*, Oospira (O.): Ninh Binh, Hoa Binh, Thanh Hoa, Nghe An.
moirati (Bavay & Dautzenberg 1909), Synprosphyma (S.) (= rudis A. J. Wagner 1922): Lao Cai, Lai Chau.
n. sp.*, Oospira (O.): Son La.
oospiroides H. Nordsieck 2011*, Synprosphyma (S.): Ha Giang.
ophthalmophana (Mabille 1887)*, Hemiphaedusa (Selenophaedusa) (+ cazioti Bavay & Dautzenberg 1909*): Lang Son, Thai Nguyen, Hoa Binh.
oviformis H. Nordsieck 2011*, Oospira (O.): Hoa Binh, Thanh Hoa.
paviei (Morlet 1892)*, Phaedusa (P.): Lai Chau, Lao Cai, Son La, Ha Giang, Yen Bai, Ha Noi, Vinh Phuc, Tuyen Quang, Bac Kan, Lang Son.
phongthoensis Loosjes & Loosjes-van Bemmel 1949*, Phaedusa (P.): Lai Chau.
polydona (Mabille 1887)*, Hemiphaedusa (Notoptychia) (= gisota Bavay & Dautzenberg 1899, = platyloma Moellendorff 1901, + montana Moellendorff 1901): Lang Son, Bac Kan.
porphyrea (Moellendorff 1882), Euphaedusa (Papilliphaedusa) (theristica Mabille 1887*, = vatheleti Bavay & Dautzenberg 1899, = gereti Bavay & Dautzenberg 1900): Ha Giang, Cao Bang, Lang Son, Bac Kan, Thai Nguyen, Tuyen Quang, Hoa Binh, Son La.
porphyrostoma (Bavay & Dautzenberg 1909)*, Hemiphaedusa (Selenophaedusa) (+ regina H. Nordsieck 2011*): Lao Cai, Ha Giang.
pseudauregani (Dautzenberg & Fischer 1908)*, Liparophaedusa: Cao Bang.
pumilio H. Nordsieck 2003, Hemiphaedusa (Hemizaptyx): Lao Cai.
pyknosoma Gittenberger & Vermeulen 2001*, Oospira (Atractophaedusa): Hai Phong.
rhopaloides (Moellendorff 1901), Oospira (Atractophaedusa) (= leptospira Moellendorff 1901, = microthyra Moellendorff 1901): Quang Ninh.
ruminiformis (Mabille 1887), Liparophaedusa: Lang Son.
semipolita (Bavay & Dautzenberg 1899)*, Oospira (O.) (= flaveola Bavay & Dautzenberg 1915, + mansonensis Moellendorff 1901): Lao Cai, Lang Son, Bac Kan, Tuyen Quang..
smithi Maassen & Gittenberger 2007*, Oospira (Atractophaedusa): Hai Phong.
sorella H. Nordsieck 2003*, Phaedusa (P.): Lao Cai, Bac Kan.
stenothyra Moellendorff 1901*, Phaedusa (P.): Thanh Hoa, Ninh Binh, Hoa Binh, Hai Phong, Quang Ninh.
suilla (Bavay & Dautzenberg 1909), Synprosphyma (S.): Lao Cai, Lai Chau.
sykesi (Bavay & Dautzenberg 1899)*, Oospira (Leptocochlea) (+ letrungthongi Grego & Szekeres 2011*): Lang Son, Bac Kan, Thai Nguyen.
thatkheana (Bavay & Dautzenberg 1899)*, Hemiphaedusa (Selenophaedusa) (+ splendida H. Nordsieck 2011*): Ha Giang, Cao Bang, Lang Son, Bac Kan, Tuyen Quang.
triptyx H. Nordsieck 2011*, Oospira (O.): Son La.
vanbuensis (Bavay & Dautzenberg 1899)*, Oospira (O.) (= umbratica Szekeres 1970, = pocsi Szekeres 1969): Ninh Binh, Hoa Binh, Son La, Thanh Hoa, Nghe An..
Garnieriinae C. Boettger 1926
ardouiniana (Heude 1885)*, Grandinenia (= horrida Mabille 1887, = sajoi Szekeres 1969): Quang Ninh, Hai Phong.
bavayi (Lindholm 1924)*, Tropidauchenia (+senescens H. Nordsieck 2011*): Bac Kan.
demangei (Bavay & Dautzenberg 1909)*, Euryauchenia: Thanh Hoa.
dorri (Bavay & Dautzenberg 1899)*, Tropidauchenia: Bac Kan.
fischeri H. Nordsieck 2002, Euryauchenia (+ reticulata H. Nordsieck 2002* = phasmoides Grego & Szekeres 2011*): Tuyen Quang?, Ninh Binh, Hoa Binh, Thanh Hoa.
giardi (Fischer 1898)*, Tropidauchenia (+ citrina H. Nordsieck 2002, + brunnea H. Nordsieck 2002): Cao Bang, Lang Son, Bac Kan.
ootanii H. Nordsieck 2007, Tropidauchenia (longicollis H. Nordsieck 2011*): Ha Giang.
orientalis (Mabille 1887)*, Tropidauchenia (+ messageri Bavay & Dautzenberg 1899): Ha Noi, Lang Son.
palatalis H. Nordsieck 2011*, Tropidauchenia: Ha Giang.
panhai H. Nordsieck 2010*, Ptychauchenia: Ninh Binh, Quang Binh.
proctostoma (Mabille 1889)*, Megalauchenia (= goliath Rolle 1910, + donggiaoensis (H. Nordsieck 2002),+ forceps Loosjes & Loosjes-van Bemmel 1973*): Ha Nam, Ninh Binh, Hoa Binh, Son La, Thanh Hoa.
soror H. Nordsieck 2011*, Megalauchenia: Thanh Hoa.
tonkinensis H. Nordsieck 2010*, Neniauchenia: Ninh Binh, Quang Binh.
yongshuae H. Nordsieck 2007*, Megalauchenia: Thanh Hoa.
Zoogeographical remarks
If one compares the ranges of the species, zoogeographical subregions of northern Vietnam are recognizable as follows:
northeastern Tonkin: Ha Giang (part), Cao Bang, Lang Son, Bac Can, Thai Nguyen, Tuyen Quang;
coastal northeastern Tonkin: Quang Ninh, Hai Phong;
northwestern Tonkin: Lai Chau, Lao Cai, Ha Giang (part), Son La (N);
southern Tonkin and northernmost Annam: Ha Nam, Ninh Binh, Hoa Binh, Son La (S), Thanh Hoa, Nghe An, Quang Binh.
Most species have been collected until now only within one subregion, hardly more than 10% within two.
The remaining part of Vietnam is not considered, because its clausiliid fauna is unknown, with the exception of one species from Annam (P. cochinchinensis).
Remarks on Schileyko's (2011) list of Vietnamese Clausiliidae:
Schileyko's (2011) check-list of land snails of Vietnam contains also a list of the Vietnamese Clausiliidae (: 11-22), which is not based on a revision of the family. Therefore there are many differences to the list presented above which are commented as follows:
S. lavillei (: 14),
1. According to my revision (examination of types) the following names are synonyms:
H. ventriculus (: 12), S. callistomella (: 13), S. falcifera (: 14),
N. gisota and N. g. platyloma (: 14), A. rhopaloides leptospira and A. r. microthyra (: 16), D. duella lyteostoma (: 17), O. pocsi and O. umbratica (: 18), P. hamonvillei (: 19), and T. goliath (: 22).
S. cazioti (: 13), N. montana (: 14), O. mansonensis (: 17), and T. messageri (: 22) are evaluated as subspecies of other species.
P. theristica (: 19) is not a Phaedusa species, but a subspecies of Euphaedusa porphyrea.
2. Schileyko subdivides the genera Hemiphaedusa (: 11-14) and Oospira (: 15-18) arbitrarily into several genera, without foundation. A subdivision of these genera can only be accepted, if it is based on a revision. On the other side, the genus Tropidauchenia (: 21-22) remains undivided; the subdivision into several genera based on shell and genital morphology (Nordsieck 2007: 73-74, 149-150) has not been considered by Schileyko.
3. The classification of the genus Synprosphyma with Hemiphaedusa (H.) (: 11-12) is wrong. The separation of this group from other Phaedusinae based on genital morphology (Nordsieck 2007: 68-69) has not been considered.
4. Clausilia fargesiana sensu Bavay & Dautzenberg 1899 [non Heude] (: 15) is a synonym of Liparophaedusa auregani (Nordsieck 1998: 130). A separate species named Excussispira fargesiana does not exist in Vietnam.
5. Until now, C. bolovenica Moellendorff from Laos (: 12) is not known from Vietnam. This species has a lunellar of plicae type and is therefore classified with Oospira (Nordsieck 2002a: 86), not with Hemiphaedusa.
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References
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