taxonID	type	description	language	source
0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	description	Figs 16, 17, 18, 19, Table 7	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	diagnosis	Diagnosis. Both morphological characteristics (body very stout, skin rough with dermal ridges and tubercles, forelimbs of males strongly enlarged, with inner side of arms or fingers or chest and belly with black spines) (Fei et al. 2009) and molecular data revealed the new species to be nested within Quasipaa. Quasipaa binhi sp. nov. is distinguishable from its congeners by a combination of the following morphological characteristics: (1) SVL 76.9 – 101.1 mm in males and 88.5 – 123.4 mm in females; (2) head broader than long (HL / HW 0.96 in males, 0.96 in females); (3) vomerine teeth present; (4) external vocal sacs absent; (5) tympanum visible, round; (6) dorsum with thin and elongate ridges intermixed with small round tubercles; (7) flanks covered by oval and round tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) dorsum and dorsal surface of fore- and hindlimbs with small black spines, scattered; (11) nuptial pad absent on finger I in males (12) ventral surface of body and all fingers without spines in males; (13) eggs yellowish cream with melanic poles in females; (14) toes fully webbed to distal end of terminal phalanx; and (15) in life, dorsum dark brown and belly immaculate white, and iris dark green.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	description	Description of holotype. A large frog (SVL 101.1 mm); habitus robust with enlarged head (HL / SVL 0.40, HW / SVL 0.42); head broader than long (HL 40.6 mm, HW 42.0 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to eye than to the tip of snout (NS 8.3 mm, EN 7.4 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 15.8 mm, ED 12.2 mm); interorbital distance smaller than internarial distance and upper eyelid width (IOD 6.7 mm, IND 9.3 mm, UEW 9.9 mm); tympanum slightly visible (TD 4.4 mm) smaller than the distance from tympanum to eye (TYE 6.1 mm), ~ 36 % eye diameter; vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; external vocal sac absent. Forelimbs: arms short; upper arm length (UAL) 17.2 mm, forearm length (FAL) 45.1 mm; relative finger lengths: II <I <IV <III; fingers free of webbing; narrow dermal ridge on sides of fingers present on fingers II, III; tips of fingers swollen, not expanded; subarticular tubercles prominent, round, formula 1, 1, 2, 2; inner metatarsal tubercle oval; outer metatarsal tubercle elongate; nuptial pad absent. Hindlimbs: tibia length longer than thigh length (FeL 56.0 mm, TbL 61.0 mm), ~ 3.7 × longer than wide (TbW 16.2 mm); tips of toes swollen, round; relative length of toes: I <II <V <III <IV; toes fully webbed to distal end of terminal phalanx; dermal ridge present on outer sides of toes I and V; subarticular tubercles prominent, elongate, formula 1, 1, 2, 3, 2; inner metatarsal tubercle elongate; outer metatarsal tubercle absent; tibio-tarsal articulation reaching to tip of snout. Skin texture in life: dorsal surface of head with oval and round tubercles, dorsum with thin and elongate ridges intermixed with small round tubercles; flanks covered by oval and round tubercles; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; dorsal surface of forelimb and hindlimb with thin and elongate ridges intermixed with small tubercles; belly and ventral surface of thigh smooth. Nuptial spines: dorsum, upper flanks, upper lip, and dorsal surface of fore- and hindlimbs with small spines, scattered; ventral surface of body and fingers without spines. Coloration in life: iris dark green; dorsum and upper part of flanks dark brown; lower part of flanks yellow brown with white tubercles and black spines on top; dorsal surface of limbs yellowish brown with dark crossbars; throat and chest white with brown markings; ventral surface of limbs and belly immaculate white; toe webbing dark brown. Coloration in preservative: coloration in preservative is the same in life but somewhat faded.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	distribution	Distribution. Quasipaa binhi sp. nov. is currently known from Quang Binh (Dong Chau-Khe Nuoc Trong Nature Reserve) and Thua Thien Hue (Sao La Nature Reserve) provinces, Vietnam. Data obtained from GenBank shows that this species was also recorded from Xekong Province, Laos (Suwannapoom et al. 2021; see Discussion below).	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	etymology	Etymology. The new species is named in honor of our colleague and friend, late Assoc. Prof. Dr. Binh Van Nguyen from the Hue University, Hue City, Vietnam, in recognition of his contributions on ecological research of amphibians in Vietnam. We recommend “ Binh’s Spiny Frog ” as the common English name of the new species and the common name in Vietnamese as “ Ếch gai s ần bình ”.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	description	Figs 7, 8, 9, 10, 11, Table 6	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	diagnosis	Diagnosis. Both morphological characteristics (body very stout, skin rough with dermal ridges and tubercles, forelimbs of males strongly enlarged, with inner side of arms, fingers or chest and belly with black spines) (Fei et al. 2009) and molecular data revealed the new species to be nested within Quasipaa. Quasipaa ohlerae sp. nov. is distinguishable from its congeners by a combination of the following morphological characteristics: (1) SVL 86.7 – 107.8 mm in males and 92.7 – 107.0 mm in females; (2) head broader than long (HL / HW 0.89 in males, 0.88 in females); (3) vomerine teeth present; (4) external vocal sacs absent; (5) tympanum visible, round; (6) dorsum with thick ridges and small round tubercles; (7) flanks covered by oval and round tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) ventral surface of arms and all fingers with spines in males; (11) fingers I and II with nuptial pad in males; (12) each chest tubercle with one black spine in males; (13) females with yellowish cream eggs; (14) toes fully webbed to distal end of terminal phalanx; and (15) in life, dorsum dark brown, chest and belly immaculate white, iris dark green.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	description	Description of holotype. A large frog (SVL 103.1 mm); habitus robust with enlarged head (HL / SVL 0.38, HW / SVL 0.43); head broader than long (HL 39.4 mm, HW 44.6 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to eye than to the tip of snout (NS 9.0 mm, EN 7.9 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 16.0 mm, ED 13.0 mm); internarial distance wider than interorbital distance and upper eyelid width (IND 10.2 mm, IOD 7.4 mm, UEW 9.3 mm); tympanum visible (TD 4.0 mm) smaller than the distance from tympanum to eye (TYE 7.0 mm), ~ 30 % eye diameter; vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; external vocal sac absent. Forelimbs: arms short; upper arm length (UAL) 23.1 mm, forearm length (FAL) 51.0 mm; relative finger lengths: II <I <IV <III; fingers free of webbing; narrow dermal ridge on sides of fingers present on fingers II, III; tips of fingers swollen, not expanded; subarticular tubercles prominent, round, formula 1, 1, 2, 2; inner metatarsal tubercle oval; outer metatarsal tubercle elongate; fingers I and II with nuptial pad. Hindlimbs: tibia length longer than thigh length (FeL 52.8 mm, TbL 60.3 mm), ~ 3.2 × longer than wide (TbW 19.1 mm); tips of toes swollen, slightly round; relative length of toes: I <II <V <III <IV; toes fully webbed to distal end of terminal phalanx; dermal ridge present on outer sides of toes I and V; subarticular tubercles prominent, elongate, formula 1, 1, 2, 3, 2; inner metatarsal tubercle elongate; outer metatarsal tubercle absent; tibio-tarsal articulation reaching to nostril. Skin texture in life: dorsal surface of head with oval and round tubercles, dorsum with thick ridges intermixed with small round tubercles; flanks covered by oval and round tubercles; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; dorsal surface of forelimbs and hindlimbs with small tubercles; belly and ventral surface of thighs smooth. Nuptial spines: body of males with spines except for ~ 1 / 3 posterior part of belly and ventral surface of hindlimbs; dense spines on lower flanks, ventral surface of forelimbs, lower lip, throat, chest, 2 / 3 anterior part of belly and fingers I, II, III; dorsum, upper flanks, upper lip, dorsal surface of fore- and hindlimbs, and finger IV with small spines, scattered; each chest tubercle with one black spine. Coloration in life: iris dark green; dorsum and upper part of flanks dark brown; lower part of flanks whitish brown with white tubercles and black spines on top; dorsal surface of limbs yellowish brown with dark crossbars; throat white with brown markings; ventral surface of limbs, chest, and belly immaculate white; toe webbing dark brown. Coloration in preservative: coloration in preservative is the same in life but somewhat faded.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	distribution	Distribution. Quasipaa ohlerae sp. nov. is currently known from Son La (Copia Nature Reserve), Thanh Hoa (Xuan Lien Nature Reserve), and Nghe An (Pu Hoat Nature Reserve) provinces, Vietnam. Data obtained from GenBank shows that this species was also recorded from Yunnan Province in China; Phongsaly Province in Laos; and Nan Province in Thailand (Suwannapoom et al. 2021; see Discussion below).	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	etymology	Etymology. The new species is named in honor of our colleague and friend, Prof. Dr. Annemarie Ohler from the Département de Systématique et Evolution, Muséum National d’Histoire Naturelle, Paris, France, in recognition of her great contributions towards a better understanding of the amphibian systematics of the Indochinese region. We recommend “ Ohler’s Spiny Frog ” as the common English name of the new species and the common name in Vietnamese as “ Ếch gai s ần ohler ”.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
C9F126A2CD8257E388FF4292FE359B20.taxon	description	Figs 4, 5, 6, Table 5	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
C9F126A2CD8257E388FF4292FE359B20.taxon	description	Description. A large frog (SVL up to 106 mm in males and 95 mm in females); habitus robust with enlarged head (HL / SVL 0.38 ± 0.01, HW / SVL 0.43 ± 0.01, in males and HL / SVL 0.38 ± 0.01, HW / SVL 0.43 ± 0.01, in females); head broader than long (HL 38.0 ± 2.8 mm, HW 42.9 ± 3.0 mm, in males and HL 32.8 ± 2.0 mm, HW 37.0 ± 2.2 mm, in females); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to eye than to the tip of snout; canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter; internarial distance wider than interorbital distance and upper eyelid width; tympanum slightly visible (TD 5.4 ± 0.5 mm, in males and 5.0 ± 0.5 mm, in females) smaller than the distance from tympanum to eye (TYE 6.3 ± 0.6 mm, in males and 5.6 ± 0.7 mm, in females), ~ 50 % eye diameter; vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; external vocal sac absent. Forelimbs: arms short; upper arm length (UAL 20.1 ± 2.8 mm, in males and UAL 15.5 ± 1.4 mm, in females), forearm length (FAL 48.5 ± 3.5 mm, in males and FAL 38.4 ± 2.6 mm, in females); relative finger lengths: II <I <IV <III; fingers free of webbing; sides of fingers I, II, and III with narrow dermal ridge; tips of fingers swollen, not expanded; subarticular tubercles prominent, round, formula 1, 1, 2, 2; inner metatarsal tubercle round; outer metatarsal tubercle elongate; finger I with nuptial pad in males. Hindlimbs: tibia length longer than thigh length (FeL 52.0 ± 3.1 mm, TbL 53.1 ± 3.0 mm, in males and FeL 44.8 ± 3.3 mm, TbL 45.7 ± 3.1 mm, in females), ~ 3 × longer than wide (TbW 18.9 ± 1.8 mm, in males and TbW 15.0 ± 1.7 mm, in females); tips of toes swollen, round; relative length of toes: I <II <V <III <IV; toes fully webbed to distal end of terminal phalanx; dermal ridge present on outer sides of toes I and V; subarticular tubercles prominent, oval, formula 1, 1, 2, 3, 2; inner metatarsal tubercle elongate; outer metatarsal tubercle absent; tibio-tarsal articulation reaching to nostril. Skin texture in life: dorsal surface of head with oval and round tubercles, dorsum with thick ridges intermixed with small round tubercles; flanks covered by oval and round tubercles; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral fold absent; dorsal surface of forelimb and hindlimb with small tubercles; belly and ventral surface of thigh smooth. Nuptial spines: body of males with spines; dense spines on lower flanks, chest, 2 / 3 anterior part of belly and fingers I, II, III; spines present on dorsum, upper flanks, upper lip, dorsal surface of fore- and hindlimbs, lower lip, and throat small and scattered; spines absent on finger IV and ventral surface of forelimbs. Coloration in life: iris pale copper; dorsum and upper part of flanks yellowish grey or pale brownish grey; lower part of flanks whitish yellow with white tubercles and black spines on top; dorsal surface of limbs yellowish with brown crossbars; ventral surface of limbs yellowish white; throat white with brown markings; chest and belly pale yellowish white; toe webbing pale brown.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
C9F126A2CD8257E388FF4292FE359B20.taxon	distribution	Distribution. The species was recorded in Lao Cai (Hoang Lien National Park), Vinh Phuc (Tam Dao National Park), Ha Giang (Bac Me Nature Reserve), and Tuyen Quang (Na Hang Nature Reserve and Cham Chu Nature Reserve) provinces, northern Vietnam.	en	Pham, Cuong The, Hoang, Chung Van, Phan, Tien Quang, Pham, Anh Van, Ong, An Vinh, Nguyen, Vien Hong Thi, Ziegler, Thomas, Nguyen, Truong Quang (2025): Taxonomic revision of the Quasipaa verrucospinosa complex (Amphibia, Dicroglossidae) in Vietnam, with descriptions of two new species. ZooKeys 1240: 139-175, DOI: 10.3897/zookeys.1240.147337
