taxonID	type	description	language	source
03E37055FFC2FF86F971FB70C876E2D5.taxon	type_taxon	Type species. Helix volvulus Müller, 1774 [51] (by original designation)	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFC2FF98F971FACDCEBDE0B4.taxon	description	Conchological description. Medium to large-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 21.1 to 34.6 mm (mean = 30.0 mm), shell breadth 25.8 to 43.1 mm (mean = 36.1 mm) (sub-group I: shell height 31.6 to 34.6 mm (mean = 33.2 mm), shell breadth 36.9 to 43.1 mm (mean = 40.5 mm); sub-group II: shell height 21.1 to 26.7 mm (mean = 24.1 mm), shell breadth 25.8 to 31.4 mm (mean = 28.1 mm )). Number of whorls 4.75 to 5.5 (sub-group I: 5.25 to 5.5; sub-group II: 4.75 to 5.25). Early whorls without bluish colouration. Shape of body whorl usually slightly compressed or uncompressed; usually rounded or sub-angular (sub-group I: usually slightly compressed or uncompressed; usually rounded; sub-group II: slightly compressed or uncompressed; rounded or sub-angular). Lip usually reflected and simple (sub-group I: reflected and simple; sub-group II: reflected and simple, or unreflected and duplicated). Lip colour ivory, white, orange, peach or yellow (sub-group I: ivory, orange, peach, white or yellow; sub-group II: ivory or white). Wing-shaped projection at columellar margin absent. Peripheral band usually dark brown (sub-group I: usually dark brown; sub-group II: dark brown). Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes, spiral lines and spiral bands in different shades of brown or in white (sub-group I: zig-zag stripes, spiral bands and spiral lines in different shades of brown or in white; sub-group II: zig-zag stripes and spiral lines in different shades of brown). Typical elements of pattern below position of peripheral band: spiral lines, spiral bands and flames in different shades of brown (sub-group I: spiral lines, spiral bands and flames in darker shades of brown; sub-group II: spiral lines, spiral bands and flames in different shades of brown).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFC2FF98F971FACDCEBDE0B4.taxon	materials_examined	Displayed specimens. VNM 019 (Fig 5 a; sub-group I), specimen in ethanol; NHMUK 20110591 (ethanol-preserved frozen tissue samples NHMUK Barcode 014041333, NHMUK Barcode 014041350); GenBank accession numbers MN 153306 (COI), MN 153355 (16 S), MN 153403 (28 S); sex of individual not determined (parts of soft body damaged); shell height 32.2 mm, shell breadth 39.0 mm, number of whorls 5.25. VNM 022 (Fig 5 b; sub-group II), specimen in ethanol; NHMUK 20110540 / 2; GenBank accession numbers MN 153309 (COI), MN 153358 (16 S); female individual; shell height 24.8 mm, shell breadth 29.0 mm, number of whorls 5.25. VNM 021 (Fig 5 c; sub-group II), specimen in ethanol; NHMUK 20110540 / 1 (ethanol-preserved frozen tissue samples NHMUK Barcode 014041331, NHMUK Barcode 014041348); GenBank accession numbers MN 153308 (COI), MN 153357 (16 S), MN 153405 (28 S); sex of individual not determined (parts of soft body damaged); shell height 26.7 mm, shell breadth 31.4 mm, number of whorls 5.25. VNM 023 (Fig 5 d; sub-group II), specimen in ethanol; NHMUK 20110541 (ethanol-preserved frozen tissue samples NHMUK Barcode 014041332, NHMUK Barcode 014041349); GenBank accession numbers MN 153310 (COI), MN 153359 (16 S), MN 153406 (28 S); sex of individual not determined (parts of soft body damaged); shell height 22.6 mm, shell breadth 26.3 mm, number of whorls 4.75. Locality and sampling data. VNM 019: Vietnam, Lang Son, Trung Quan commune (22.13137 ˚, 106.58715 ˚); 23 May 2011; leg. Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 022, VNM 021, VNM 023: Vietnam, Bac Kan, Bang Van commune (22.47878 ˚, 106.04597 ˚); 19 May 2011; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFC2FF98F971FACDCEBDE0B4.taxon	diagnosis	Diagnosis. Cyclophorus courbeti can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 10: C, 25: C, 40: G, 127: A, 184: G, 214: A, 292: C, 294: C, 436: A, 493: T, 508: C, 514: G, 604: A, 638: A, 652: C, 653: C (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFC2FF98F971FACDCEBDE0B4.taxon	discussion	Remarks. Despite extensive effort, we were unable to locate type material of this species (see also [107]). Our determination was thus based on the description and figure in Ancey [92]. The description and the depicted shell matched sub-group I in size, shape (but the studied material of sub-group I had a less pronounced keel, which was, however, present in sub-group II) and colouration. The type locality given in Ancey [92] is “ upper Tonkin, in the region of Lang Son and Bac Ninh ” (original: “ Haut-Tonkin, dans la région de Lang-son et de Bac-Ninh ”), and our studied samples of sub-group I were collected from a locality in this area. We regard the material from eastern Thailand, which has been determined as C. courbeti by Nantarat et al. [47] (also included in the phylogenies of [6] and this study, herein THA 011), as belonging to a different species. The studied material of C. courbeti has been found in northern Lang Son province (sub-group I) and in northeastern Bac Kan province (sub-group II) (Fig 4). Another herein characterised species, C. subfloridus (sub-group II), occurs sympatrically with C. courbeti (sub-group I) in northern Lang Son province. The shell shape of C. subfloridus is generally less compressed than that of C. courbeti (Fig 2). The shell of C. subfloridus sub-group II is smaller than that of C. courbeti sub-group I (Fig 3 b) and the early whorls show a bluish colouration, while those of C. courbeti sub-group I lack a bluish colouration. The lip of C. subfloridus sub-group II tends to have a more whitish colour than that of C. courbeti sub-group I, which tends to be more yellowish or reddish.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDCFF9BF971F96CC900E46E.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 23.8 to 33.3 mm (mean = 29.0 mm), shell breadth 29.6 to 39.4 mm (mean = 34.2 mm) (sub-group I: shell height 26.2 to 33.3 mm (mean = 29.9 mm), shell breadth 32.5 to 39.4 mm (mean = 35.8 mm); sub-group II: shell height 23.8 to 32.2 mm (mean = 28.1 mm), shell breadth 29.6 to 36.0 mm (mean = 32.5 mm )). Number of whorls 4.75 to 5.25 (sub-group I: 4.75 to 5.25; sub-group II: 5 to 5.25). Early whorls usually without bluish colouration (sub-group I: usually without bluish colouration; sub-group II: without bluish colouration). Shape of body whorl usually uncompressed or slightly compressed; rounded, sub-angular or well-rounded (sub-group I: usually uncompressed or slightly compressed; rounded or sub-angular; sub-group II: usually uncompressed; rounded or well-rounded). Lip reflected and simple. Lip colour red, yellow, ivory, orange or white (sub-group I: yellow, ivory, orange or white; sub-group II: red). Wing-shaped projection at columellar margin usually absent (sub-group I: absent, or present and not pronounced; sub-group II: absent). Peripheral band usually dark brown (sub-group I: dark brown; sub-group II: usually dark brown). Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes, spiral bands, spiral lines and flames in different shades of brown or in white (sub-group I: zig-zag stripes, spiral bands, spiral lines and flames in different shades of brown or in white; sub-group II: zig-zag stripes, spiral bands, spiral lines and flames in different shades of brown). Typical elements of pattern below position of peripheral band: spiral lines, spiral bands and flames in different shades of brown (sub-group I: spiral lines, spiral bands and flames in darker shades of brown; sub-group II: spiral lines, spiral bands and flames in different shades of brown).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDCFF9BF971F96CC900E46E.taxon	materials_examined	Displayed specimens. VNM 005 (Fig 6 a; sub-group I), specimen in ethanol; NHMUK 20110513 / 3; GenBank accession numbers MG 720873 (COI), MG 720923 (16 S), MG 720973 (28 S); female individual; shell height 30.0 mm, shell breadth 35.5 mm, number of whorls 5.25. VNM 007 (Fig 6 b; sub-group I), specimen in ethanol; NHMUK 20110513 / 2 (ethanol-preserved frozen tissue samples NHMUK Barcode 014041330, NHMUK Barcode 014041347); GenBank accession numbers MN 153301 (COI), MN 153350 (16 S), MN 153399 (28 S); female individual; shell height 31.3 mm, shell breadth 36.1 mm, number of whorls 5.25. VNM 008 (Fig 6 c; sub-group I), specimen in ethanol; NHMUK 20110513 / 4; GenBank accession numbers MG 720874 (COI), MG 720924 (16 S), MG 720974 (28 S); male individual; shell height 28.0 mm, shell breadth 32.5 mm, number of whorls 5.25. VNM 015 (Fig 6 d; sub-group II), specimen in ethanol; NHMUK 20110476 / 9; GenBank accession numbers MG 720865 (COI), MG 720915 (16 S), MG 720965 (28 S); male individual; shell height 28.4 mm, shell breadth 31.6 mm, number of whorls 5.25. VNM 013 (Fig 6 e; sub-group II), specimen in ethanol; NHMUK 20110476 / 7; GenBank accession numbers MG 720863 (COI), MG 720913 (16 S), MG 720963 (28 S); female individual; shell height 27.9 mm, shell breadth 33.2 mm, number of whorls 5. VNM 014 (Fig 6 f; sub-group II), specimen in ethanol; NHMUK 20110476 / 8; GenBank accession numbers MG 720864 (COI), MG 720914 (16 S), MG 720964 (28 S); male individual; shell height 26.3 mm, shell breadth 30.0 mm, number of whorls 5. Locality and sampling data. VNM 005, VNM 007, VNM 008: Vietnam, Bac Kan, Ba Be National Park (22.40923 ˚, 105.62855 ˚); 18 May 2011; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 015, VNM 013, VNM 014: Vietnam, Thai Nguyen, Dong Dat commune (21.75523 ˚, 105.70745 ˚); 16 May 2011; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDCFF9BF971F96CC900E46E.taxon	diagnosis	Diagnosis. Cyclophorus fasciatus can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 127: G, 184: A, 214: G / C, 232: C, 271: C / T, 313: A, 436: G, 493: C, 571: C, 604: G (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDCFF9BF971F96CC900E46E.taxon	discussion	Remarks. This species was originally described as Cyclophorus dodrans fasciatus. The holotype (SMF 34732) resembled sub-group I in size (but the holotype was slightly larger), shape and colouration (but not in the exact colour pattern). The type locality given in Kobelt [48], “ Residence in Annam ” (original: “ Aufenthalt in Annam ”), is most likely incorrect as Zilch [108] states that “ inside the shell, a locality label was found: Tuyen Quang, Tonkin. Fruhstorfer ” (original: “ im Inneren des Gehäuses fand sich ein Fundortzettel: Tuyen-Quang, Tonkin. Fruhstorfer ”). Our studied samples of sub-group I were partly collected very close to northeastern Tuyen Quang. Further material from Tuyen Quang should be studied in future to confirm our determination. This species corresponds to lineage 03 in Oheimb et al. [6]. The studied material of C. fasciatus has been found in Ba Be National Park in northwestern Bac Kan province (sub-group I), in eastern Thai Nguyen province (sub-group I) and in central Thai Nguyen province (sub-group II) (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species. Cyclophorus subfloridus (sub-group I) occurs in southwestern Lang Son province, approx. 20 km from where C. fasciatus (sub-group I) was found, in eastern Thai Nguyen province. The shell shape of C. subfloridus is generally slightly less compressed than that of C. fasciatus (Fig 2). The shell of C. subfloridus sub-group I is slightly smaller than that of C. fasciatus sub-group I (Fig 3 b) and the early whorls show a bluish colouration, while those of C. fasciatus sub-group I usually lack a bluish colouration. The lip of C. subfloridus sub-group I tends to have a more whitish colour than that of C. fasciatus sub-group I, which tends to be more yellowish or reddish.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDFFF9DF971FD82C882E4DC.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 21.8 to 31.1 mm (mean = 27.6 mm), shell breadth 25.1 to 34.0 mm (mean = 30.6 mm) (sub-group I: shell height 26.0 to 31.1 mm (mean = 28.4 mm), shell breadth 28.0 to 34.0 mm (mean = 31.3 mm); sub-group II: shell height 21.8 to 24.8 mm (mean = 23.0 mm), shell breadth 25.1 to 29.1 mm (mean = 26.8 mm )). Number of whorls 5 to 5.5 (sub-group I: 5.25 to 5.5; sub-group II: 5). Early whorls with or without bluish colouration (sub-group I: with or without bluish colouration; sub-group II: with bluish colouration). Shape of body whorl usually uncompressed or slightly compressed; well-rounded (sub-group I: uncompressed or slightly compressed; well-rounded; sub-group II: uncompressed, slightly compressed or compressed; well-rounded). Lip usually reflected and simple (sub-group I: usually reflected and simple; sub-group II: reflected and simple). Lip colour white or ivory (sub-group I: ivory or white; sub-group II: white). Wing-shaped projection at columellar margin present and pronounced. Peripheral band dark brown. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: flames, zig-zag stripes and spots in different shades of brown or in white (sub-group I: flames, zig-zag stripes and spots in different shades of brown or in white; sub-group II: flames and zig-zag stripes in darker shades of brown). Typical elements of pattern below position of peripheral band: spiral lines, spiral bands and flames in darker shades of brown (sub-group I: spiral lines, spiral bands and flames in darker shades of brown; sub-group II: spiral lines and flames in darker shades of brown).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDFFF9DF971FD82C882E4DC.taxon	materials_examined	Displayed specimens. VNM 042 (Fig 7 a; sub-group I), specimen in ethanol; NHMUK 20110608 / 3; GenBank accession numbers MG 720878 (COI), MG 720928 (16 S), MG 720978 (28 S); male individual; shell height 27.0 mm, shell breadth 29.8 mm, number of whorls 5.25. VNM 043 (Fig 7 b; sub-group I), specimen in ethanol; NHMUK 20110608 / 4; GenBank accession numbers MN 153319 (COI), MN 153368 (16 S), MN 153415 (28 S); female individual; shell height 28.1 mm, shell breadth 29.7 mm, number of whorls 5.5. VNM 044 (Fig 7 c; sub-group I), specimen in ethanol; NHMUK 20110608 / 2 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041336); GenBank accession numbers MG 720877 (COI), MG 720927 (16 S), MG 720977 (28 S); female individual; shell height 26.5 mm, shell breadth 29.6 mm, number of whorls 5.25. VNM 045 (Fig 7 d; sub-group II), specimen in ethanol; NHMUK 20110603 / 2; GenBank accession numbers MG 720876 (COI), MG 720926 (16 S), MG 720976 (28 S); male individual; shell height 21.8 mm, shell breadth 25.1 mm, number of whorls 5. VNM 046 (Fig 7 e; sub-group II), specimen in ethanol; NHMUK 20110603 / 1 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041335); GenBank accession numbers MN 153320 (COI), MN 153369 (16 S), MN 153416 (28 S); sex of individual not determined (parts of soft body damaged); shell height 24.8 mm, shell breadth 29.1 mm, number of whorls 5. Locality and sampling data. VNM 042, VNM 043, VNM 044: Vietnam, Quang Ninh, Ha Long, Ha Phong ward (20.96638 ˚, 107.16610 ˚); 25 May 2011; leg. Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 045, VNM 046: Vietnam, Quang Ninh, Son Duong commune (21.06693 ˚, 106.98883 ˚); 24 May 2011; leg. Hao Van Luong.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDFFF9DF971FD82C882E4DC.taxon	diagnosis	Diagnosis. Cyclophorus jourdyi can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 184: C, 325: C, 415: T, 458: G (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDFFF9DF971FD82C882E4DC.taxon	discussion	Remarks. The studied syntypes (MNHN-IM- 2000 - 33830, MNHN-IM- 2000 - 33831 and RBINS I. G. 10591 / MT. 902, the latter labelled as paratype) resembled sub-group I and II in size, shape and colouration. No specific type locality is given in Morlet [93] besides “ Tonkin ”, where all taxa from the respective publication have been described from. In Morlet [109], however, the following locality is given for this species: “ Tonkin: Ha Long Bay and Elephant Mountain ” (original: “ Tonkin: baie d’Halong et montagne de l’Éléphant ”), which includes “ Ha Long Bay / Tonkin ” (original: “ Baie d’Halong / Tonkin ”) given on a label of the Brussels syntype (the Paris syntypes’ labels only state “ Tonkin ” (original: “ Tonkin ” and “ Tonquin ” )) and the location where our studied samples of sub-group I were collected. This species corresponds to lineage 07 in Oheimb et al. [6]. We regard the material from central Vietnam, which has been determined as C. jourdyi by Nantarat et al. [47] (also included in the phylogenies of [6] and this study, herein VNM 029), as belonging to a different species. The studied material of C. jourdyi has been found in southern Quang Ninh province (sub-groups I and II) (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD9FF9FF971FD34C9EFE021.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 22.3 to 30.4 mm (mean = 26.7 mm), shell breadth 23.9 to 33.7 mm (mean = 29.6 mm) (sub-group I: shell height 25.2 to 30.4 mm (mean = 28.2 mm), shell breadth 29.0 to 33.7 mm (mean = 31.4 mm); sub-group II: shell height 22.3 to 27.4 mm (mean = 25.2 mm), shell breadth 23.9 to 30.3 mm (mean = 27.7 mm )). Number of whorls 5 to 5.5 (sub-group I: 5.25 to 5.5; sub-group II: 5 to 5.25). Early whorls with bluish colouration. Shape of body whorl usually uncompressed; rounded or well-rounded (sub-group I: usually uncompressed; well-rounded or rounded; sub-group II: uncompressed; usually rounded). Lip reflected and simple, or unreflected and duplicated (sub-group I: reflected and simple; sub-group II: reflected and simple, or unreflected and duplicated). Lip colour ivory or white. Wing-shaped projection at columellar margin absent, present and not pronounced, or present and pronounced (sub-group I: present and not pronounced, present and pronounced, or absent; sub-group II: absent, or present and not pronounced). Peripheral band usually dark brown (sub-group I: dark brown; sub-group II: usually dark brown). Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes and flames in different shades of brown or in white (sub-group I: zig-zag stripes and flames in different shades of brown; sub-group II: zig-zag stripes and flames in different shades of brown or in white). Typical elements of pattern below position of peripheral band: spiral lines, spiral bands, zig-zag stripes and transverse lines in different shades of brown or in white (sub-group I: spiral lines, spiral bands and transverse lines in darker shades of brown; sub-group II: zig-zag stripes, spiral bands and spiral lines in different shades of brown or in white). Displayed specimens. VNM 063 (Fig 8 a; sub-group I), specimen in ethanol; NHMUK 20140597 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041340); GenBank accession numbers MN 153334 (COI), MN 153383 (16 S), MN 153430 (28 S); female individual; shell height 27.6 mm, shell breadth 31.0 mm, number of whorls 5.25. VNM 059 (Fig 8 b; sub-group I), specimen in ethanol; NHMUK 20140557 / 2; GenBank accession numbers MN 153330 (COI), MN 153379 (16 S), MN 153426 (28 S); male individual; shell height 28.2 mm, shell breadth 31.1 mm, number of whorls 5.25. VNM 061 (Fig 8 c; sub-group I), specimen in ethanol; NHMUK 20140585 / 1 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041339); GenBank accession numbers MN 153332 (COI), MN 153381 (16 S), MN 153428 (28 S); female individual; shell height 28.4 mm, shell breadth 31.5 mm, number of whorls 5.25. VNM 065 (Fig 8 d; sub-group II), specimen in ethanol; NHMUK 20110594 (ethanol-preserved frozen tissue samples NHMUK Barcode 014041334, NHMUK Barcode 014041351); GenBank accession numbers MN 153336 (COI), MN 153385 (16 S), MN 153432 (28 S); sex of individual not determined (parts of soft body damaged); shell height 24.7 mm, shell breadth 26.8 mm, number of whorls 5.25. Locality and sampling data. VNM 063: Vietnam, Lang Son, Yen Thinh commune (21.61862 ˚, 106.34760 ˚); 19 September 2013; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 059: Vietnam, Lang Son, Huu Lien commune, Huu Lien Nature Reserve (21.66235 ˚ – 21.65998 ˚, 106.36633 ˚ – 106.36462 ˚); 17 September 2013; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 061: Vietnam, Lang Son, Yen Thinh commune (21.65948 ˚, 106.36535 ˚); 18 September 2013; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 065: Vietnam, Lang Son, Trung Quan commune (22.13137 ˚, 106.58715 ˚); 23 May 2011; leg. Hao Van Luong, Fred Naggs & Sang Van Pham.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD9FF9FF971FD34C9EFE021.taxon	diagnosis	Diagnosis. Cyclophorus subfloridus can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 217: A (number indicates the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD9FF9FF971FD34C9EFE021.taxon	discussion	Remarks. This species was originally described as Cyclophorus fulguratus var. subflorida. Despite extensive effort, we were unable to locate type material of this species (see also [107]). Our determination was thus based on the description and figure in Ancey [92]. The description and the depicted shell matched sub-group I in size, shape and colouration. The material of C. fulguratus studied in Ancey [92], which includes the therein newly described C. fulguratus var. subflorida (a specific type locality for this taxon is not given), is “ from Lang Son to Bac Ninh ” (original: “ De Lang-son a Bac-Ninh ”), and our studied samples of sub-group I were collected from localities in this area. We regard Cyclophorus fulguratus var. barniana Ancey, 1888 [92] as a synonym of this species given that the type material of both taxa was presumably collected in the same region [92] and that the described morphological differences between them are insignificant (note that no type material of C. fulguratus var. barniana could be located; see also [107]). Cyclophorus jourdyi var. longsonensis Morlet, 1891 [110] is regarded as a further synonym of C. subfloridus given that its type locality is “ Lang Son ” (original: “ Long-son ”) and that the studied type material (RBINS I. G. 10591 / MT. 903, labelled as paratype, and MNHNIM- 2000 - 34483 and MNHN-IM- 2000 - 34484, both potential syntypes) resembled sub-group I of C. subfloridus in size, shape and colouration (only in MNHN-IM- 2000 - 34483 the early whorls lacked a bluish colouration). We regard the material from northern Thailand, which has been determined as C. subfloridus by Nantarat et al. [47] (also included in the phylogenies of [6] and this study, herein THA 025), as belonging to a different species. The studied material of C. subfloridus has been found in southwestern Lang Son province, partly in Huu Lien Nature Reserve, (sub-group I) and in northern Lang Son province (sub-group II) (Fig 4). Another herein characterised species, C. courbeti (sub-group I), occurs sympatrically with C. subfloridus (sub-group II) in northern Lang Son province. The shell shape of C. courbeti is generally more compressed than that of C. subfloridus (Fig 2). The shell of C. courbeti sub-group I is larger than that of C. subfloridus sub-group II (Fig 3 b) and the early whorls lack a bluish colouration, while those of C. subfloridus sub-group II show a bluish colouration. The lip of C. courbeti sub-group I tends to have a more yellowish or reddish colour than that of C. subfloridus sub-group II, which tends to be more whitish. Furthermore, C. fasciatus (sub-group I) occurs in eastern Thai Nguyen province, approx. 20 km from where C. subfloridus (sub-group I) was found, in southwestern Lang Son province. The shell shape of C. fasciatus is generally slightly more compressed than that of C. subfloridus (Fig 2). The shell of C. fasciatus sub-group I is slightly larger than that of C. subfloridus sub-group I (Fig 3 b) and the early whorls usually lack a bluish colouration, while those of C. subfloridus sub-group I show a bluish colouration. The lip of C. fasciatus sub-group I tends to have a more yellowish or reddish colour than that of C. subfloridus sub-group I, which tends to be more whitish.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	description	urn: lsid: zoobank. org: act: 0 E 86 BA 4 B- 769 A- 4 D 8 D-AD 00 - ACA 4 EAEC 82 D 6	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	etymology	Etymology. The name refers to the type locality of this species in Cuc Phuong National Park.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 23.3 to 29.9 mm (mean = 26.5 mm), shell breadth 28.7 to 35.4 mm (mean = 31.4 mm). Number of whorls 5 to 5.25. Early whorls without or with bluish colouration. Shape of body whorl usually uncompressed or slightly compressed; rounded or well-rounded. Lip reflected and simple. Lip colour yellow or ivory. Wing-shaped projection at columellar margin present and pronounced, or present and not pronounced. Peripheral band usually dark brown, medium brown, or not distinguishable from background colouration. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: flames and zig-zag stripes in different shades of brown. Typical elements of pattern below position of peripheral band: spiral lines and spiral bands in different shades of brown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	materials_examined	Type specimens. VNM 052 (Fig 9 a; holotype), specimen in ethanol; NHMUK 20140276 / 4; GenBank accession numbers MG 720887 (COI), MG 720937 (16 S), MG 720987 (28 S); female individual; shell height 27.2 mm, shell breadth 32.7 mm, number of whorls 5.25. VNM 053 (Fig 9 b; paratype), specimen in ethanol; NHMUK 20140287 / 2 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041337); GenBank accession numbers MN 153325 (COI), MN 153374 (16 S), MN 153421 (28 S); male individual; shell height 27.0 mm, shell breadth 31.3 mm, number of whorls 5.25. VNM 054 (Fig 9 c; paratype), specimen in ethanol; NHMUK 20140287 / 3 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041338; viable cell preparation NHMUK Barcode 014041352); GenBank accession numbers MN 153326 (COI), MN 153375 (16 S), MN 153422 (28 S); male individual; shell height 27.0 mm, shell breadth 31.1 mm, number of whorls 5.25. Locality and sampling data. VNM 052: Vietnam, Ninh Binh, Cuc Phuong National Park (20.28845 ˚, 105.66767 ˚); 07 September 2013; leg. Hao Van Luong & team. VNM 053, VNM 054: Vietnam, Ninh Binh, Cuc Phuong National Park (20.35548 ˚, 105.63342 ˚); 08 September 2013; leg. Hao Van Luong & team.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	diagnosis	Diagnosis. Cyclophorus cucphuongensis sp. nov. can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 89: T, 91: C, 208: T, 289: T, 343: C, 401: C, 487: C, 556: G, 574: G / C (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFDBFF91F971F9D9CE56E574.taxon	discussion	Remarks. This species corresponds to lineage 10 in Oheimb et al. [6]. The studied material of C. cucphuongensis has been found in central and eastern Cuc Phuong National Park in both western Ninh Binh province and southeastern Hoa Binh province (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species. Cyclophorus takumisaitoi occurs in northwestern Cuc Phuong National Park. The shell shape of C. takumisaitoi is similar to that of C. cucphuongensis (Fig 2). The shell of C. takumisaitoi is slightly smaller than that of C. cucphuongensis (Fig 3 a). The lip of C. takumisaitoi tends to have a more reddish colour than that of C. cucphuongensis, which tends to be more yellowish. The wing-shaped projection at the columellar margin is absent in C. takumisaitoi, while being present in C. cucphuongensis, and the shell colouration of C. takumisaitoi is more uniform with a less prominent peripheral band. Furthermore, C. paracucphuongensis occurs in western Cuc Phuong National Park. The shell shape of C. paracucphuongensis is similar to that of C. cucphuongensis (Fig 2). The shell of C. paracucphuongensis is of similar size as that of C. cucphuongensis (Fig 3 a). On the basis of currently available shell morphological data, these two species cannot be differentiated.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	description	urn: lsid: zoobank. org: act: 803 C 12 C 9 - D 401 - 49 C 7 - 835 E- 3922 EF 9873 B 4	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	etymology	Etymology. The name refers to C. cucphuongensis, which is morphologically similar to this species.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 24.4 to 26.7 mm (mean = 25.8 mm), shell breadth 28.2 to 30.9 mm (mean = 29.8 mm). Number of whorls 5 to 5.25. Early whorls without bluish colouration. Shape of body whorl uncompressed; rounded or sub-angular. Lip reflected and simple. Lip colour yellow or ivory. Wing-shaped projection at columellar margin absent, present and not pronounced, or present and pronounced. Peripheral band dark brown. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes and flames in different shades of brown. Typical elements of pattern below position of peripheral band: spiral bands and spiral lines in darker shades of brown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	materials_examined	Type specimens. VNM 069 (Fig 10 a; holotype), specimen in ethanol; NHMUK 20180432; GenBank accession numbers MN 153337 (COI), MN 153386 (16 S), MN 153433 (28 S); male individual; shell height 25.6 mm, shell breadth 28.6 mm, number of whorls 5.25. VNM 071 (Fig 10 b; paratype), specimen in ethanol; NHMUK 20180433; GenBank accession numbers MN 153339 (COI), MN 153388 (16 S), MN 153435 (28 S); male individual; shell height 26.4 mm, shell breadth 30.3 mm, number of whorls approx. 5 (pieces of first whorl missing). VNM 070 (Fig 10 c; paratype), specimen in ethanol; NHMUK 20180434; GenBank accession numbers MN 153338 (COI), MN 153387 (16 S), MN 153434 (28 S); female individual; shell height 26.1 mm, shell breadth 30.9 mm, number of whorls 5.25. Locality and sampling data. VNM 069, VNM 071, VNM 070: Vietnam, Thanh Hoa, Cuc Phuong National Park (20.35583 ˚, 105.51088 ˚); 02 May 2009; leg. Hao Van Luong.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	diagnosis	Diagnosis. Cyclophorus paracucphuongensis sp. nov. can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 38: A, 485: C (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD5FF92F971FCACC8E2E3CC.taxon	discussion	Remarks. The studied material of C. paracucphuongensis has been found in western Cuc Phuong National Park in northern Thanh Hoa province (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species. Cyclophorus takumisaitoi occurs in northwestern Cuc Phuong National Park. The shell shape of C. takumisaitoi is similar to that of C. paracucphuongensis (Fig 2). The shell of C. takumisaitoi is slightly smaller than that of C. paracucphuongensis (Fig 3 a). The lip of C. takumisaitoi tends to have a more reddish colour than that of C. paracucphuongensis, which tends to be more yellowish. The shell colouration of C. takumisaitoi is more uniform with a less prominent peripheral band. Furthermore, C. cucphuongensis occurs in central and eastern Cuc Phuong National Park. The shell shape of C. cucphuongensis is similar to that of C. paracucphuongensis (Fig 2). The shell of C. cucphuongensis is of similar size as that of C. paracucphuongensis (Fig 3 a). On the basis of currently available shell morphological data, these two species cannot be differentiated.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	description	urn: lsid: zoobank. org: act: FD 4 FE 74 D- 974 A- 4 C 6 F-B 968 - 15 A 09 D 2 B 1 BB 0	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	etymology	Etymology. The name refers to the type locality of this species in Phong Nha-Ke Bang National Park.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 20.5 to 26.2 mm (mean = 24.1 mm), shell breadth 26.0 to 32.8 mm (mean = 29.8 mm). Number of whorls 4.75 to 5.25. Early whorls without or with bluish colouration. Shape of body whorl slightly compressed or compressed; usually rounded. Lip reflected and simple. Lip colour ivory, white or peach. Wing-shaped projection at columellar margin absent, or present and not pronounced. Peripheral band usually dark brown. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes, spiral lines and spiral bands in different shades of brown. Typical elements of pattern below position of peripheral band: spiral lines, spiral bands and transverse lines in darker shades of brown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	materials_examined	Type specimens. VNM 032 (Fig 11 a; holotype), specimen in ethanol; NHMUK 20130904 / 4; GenBank accession numbers MN 153314 (COI), MN 153363 (16 S), MN 153410 (28 S); female individual; shell height 24.5 mm, shell breadth 29.6 mm, number of whorls 5. VNM 035 (Fig 11 b; paratype), specimen in ethanol; NHMUK 20130894 / 3; GenBank accession numbers MN 153315 (COI), MN 153364 (16 S), MN 153411 (28 S); female individual; shell height 24.0 mm, shell breadth 29.4 mm, number of whorls 5. VNM 031 (Fig 11 c; paratype), specimen in ethanol; NHMUK 20130904 / 3; GenBank accession numbers MG 720883 (COI), MG 720933 (16 S), MG 720983 (28 S); male individual; shell height 23.2 mm, shell breadth 27.3 mm, number of whorls 5. Locality and sampling data. VNM 032, VNM 031: Vietnam, Quang Binh, Phong Nha-Ke Bang National Park (17.50317 ˚, 106.26045 ˚); 03 March 2012; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham. VNM 035: Vietnam, Quang Binh, Phong Nha-Ke Bang National Park (17.54147 ˚, 106.23532 ˚); 03 March 2012; leg. Jonathan Ablett, Hao Van Luong, Fred Naggs & Sang Van Pham.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	diagnosis	Diagnosis. Cyclophorus phongnhakebangensis sp. nov. can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 46: G, 95: C, 133: A, 178: C, 242: C, 259: C, 277: C, 388: G, 391: A / G, 409: G, 412: C, 415: C, 433: T, 439: A, 473: A, 474: G, 475: C, 544: G, 565: C, 595: C, 637: C (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD6FF94F971FA24CE10E2C1.taxon	discussion	Remarks. This species corresponds to lineage 06 in Oheimb et al. [6]. The studied material of C. phongnhakebangensis has been found in Phong Nha-Ke Bang National Park in central Quang Binh province (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	description	urn: lsid: zoobank. org: act: 6 D 497 FF 9 - 5849 - 49 BB- 8162 - 665 FADA 40 C 34	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	etymology	Etymology. The name refers to the malacologist and evolutionary biologist Takumi Saito, who collected individuals of this species together with Tu Van Do and Takahiro Hirano.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 22.7 to 25.0 mm (mean = 23.8 mm), shell breadth 27.1 to 29.2 mm (mean = 28.0 mm). Number of whorls 5 to 5.25. Early whorls without bluish colouration. Shape of body whorl uncompressed or slightly compressed; rounded or well-rounded. Lip reflected and simple. Lip colour orange, peach or yellow. Wing-shaped projection at columellar margin absent. Peripheral band not distinguishable from background colouration, or absent. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: zig-zag stripes and flames in lighter shades of brown. Typical elements of pattern below position of peripheral band: spiral bands in darker shades of brown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	materials_examined	Type specimens. VNM 076 (Fig 12 a; holotype), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180455 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041344); GenBank accession numbers MN 153344 (COI), MN 153393 (16 S), MN 153440 (28 S); male individual; shell height 25.0 mm, shell breadth 28.2 mm, number of whorls 5.25. VNM 078 (Fig 12 b; paratype), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180456 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041345); GenBank accession numbers MN 153346 (COI), MN 153395 (16 S), MN 153442 (28 S); male individual; shell height 23.2 mm, shell breadth 27.1 mm, number of whorls 5. VNM 077 (Fig 12 c; paratype), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180457 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041346); GenBank accession numbers MN 153345 (COI), MN 153394 (16 S), MN 153441 (28 S); male individual; shell height 23.4 mm, shell breadth 28.0 mm, number of whorls 5.25. Locality and sampling data. VNM 076, VNM 078, VNM 077: Vietnam, Hoa Binh, Cuc Phuong National Park (20.38493 ˚, 105.52280 ˚); 26 May 2016; leg. Tu Van Do, Takahiro Hirano & Takumi Saito.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	diagnosis	Diagnosis. Cyclophorus takumisaitoi sp. nov. can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 241: A, 319: C, 349: T, 430: G, 499: A (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD0FF95F971FB39C93DE29E.taxon	discussion	Remarks. The studied material of C. takumisaitoi has been found in northwestern Cuc Phuong National Park in southeastern Hoa Binh province (Fig 4). None of the other herein characterised species is known to occur sympatrically with this species. Cyclophorus cucphuongensis occurs in central and eastern Cuc Phuong National Park. The shell shape of C. cucphuongensis is similar to that of C. takumisaitoi (Fig 2). The shell of C. cucphuongensis is slightly larger than that of C. takumisaitoi (Fig 3 a). The lip of C. cucphuongensis tends to have a more yellowish colour than that of C. takumisaitoi, which tends to be more reddish. The wing-shaped projection at the columellar margin is present in C. cucphuongensis, while being absent in C. takumisaitoi, and the shell colouration of C. cucphuongensis is more vivid with a more prominent peripheral band. Furthermore, C. paracucphuongensis occurs in western Cuc Phuong National Park. The shell shape of C. paracucphuongensis is similar to that of C. takumisaitoi (Fig 2). The shell of C. paracucphuongensis is slightly larger than that of C. takumisaitoi (Fig 3 a). The lip of C. paracucphuongensis tends to have a more yellowish colour than that of C. takumisaitoi, which tends to be more reddish. The shell colouration of C. paracucphuongensis is more vivid with a more prominent peripheral band.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD1FF96F971FB72C817E0A0.taxon	description	Conchological description. Medium-sized Cyclophorus species, overall shell shape rounded, aperture typically circular. Shell height 23.1 to 26.5 mm (mean = 25.0 mm), shell breadth 28.7 to 31.9 mm (mean = 29.9 mm). Number of whorls 5.25. Early whorls with or without bluish colouration. Shape of body whorl uncompressed or slightly compressed; rounded or well-rounded. Lip reflected and simple. Lip colour yellow. Wing-shaped projection at columellar margin present and not pronounced, absent, or present and pronounced. Peripheral band dark brown or absent. Colour pattern generally variable. Typical elements of pattern above position of peripheral band: spiral lines, spiral bands and zig-zag stripes in different shades of brown. Typical elements of pattern below position of peripheral band: spiral lines, spiral bands and transverse lines in darker shades of brown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD1FF96F971FB72C817E0A0.taxon	materials_examined	Displayed specimens. VNM 073 (Fig 13 a), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180448 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041342); GenBank accession numbers MN 153341 (COI), MN 153390 (16 S), MN 153437 (28 S); male individual; shell height 24.6 mm, shell breadth 28.7 mm, number of whorls 5.25. VNM 074 (Fig 13 b), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180447 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041341); GenBank accession numbers MN 153342 (COI), MN 153391 (16 S), MN 153438 (28 S); male individual; shell height 25.5 mm, shell breadth 29.5 mm, number of whorls approx. 5.25 (pieces of first whorl missing). VNM 075 (Fig 13 c), soft body in ethanol (previously in formalin) and dry shell; NHMUK 20180452 (ethanol-preserved frozen tissue sample NHMUK Barcode 014041343); GenBank accession numbers MN 153343 (COI), MN 153392 (16 S), MN 153439 (28 S); female individual; shell height 26.5 mm, shell breadth 30.7 mm, number of whorls approx. 5.25 (pieces of first whorl missing). Locality and sampling data. VNM 073, VNM 074, VNM 075: Vietnam, Hoa Binh, Cao Duong commune, bought at market stall (20.72542 ˚, 105.64899 ˚), collected nearby according to the merchant; purchased on 26 May 2016; purchased by Tu Van Do, Takahiro Hirano & Takumi Saito.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD1FF96F971FB72C817E0A0.taxon	diagnosis	Diagnosis. Cyclophorus sp. 1 can be differentiated from all herein characterised Cyclophorus species based on DNA sequence data. Diagnostic nucleotide substitutions of the cytochrome c oxidase subunit I (COI) gene: 55: A, 94: A, 97: G, 412: G, 511: G (numbers indicate the position in the studied COI fragment).	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
03E37055FFD1FF96F971FB72C817E0A0.taxon	discussion	Remarks. This species is so far undescribed. As the studied material of Cyclophorus sp. 1 has been bought on a market, the species’ exact distribution remains unknown.	en	von Oheimb, Katharina C. M., Oheimb, Parm Viktor von, Hirano, Takahiro, Do, Tu Van, Ablett, Jonathan, Luong, Hao Van, Pham, Sang Van, Naggs, Fred (2019): Cryptic diversity of limestone karst inhabiting land snails (Cyclophorus spp.) in northern Vietnam, their evolutionary history and the description of four new species. PLoS ONE (e 0222163) 14 (10): 1-37, DOI: 10.1371/journal.pone.0222163
