identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
0C443FD8DB6F5ADB8080B11DBD3A9B6C.text	0C443FD8DB6F5ADB8080B11DBD3A9B6C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Quasipaa binhi Pham & Hoang & Phan & Pham & Ong & Nguyen & Ziegler & Nguyen 2025	<div><p>Quasipaa binhi sp. nov.</p><p>Figs 16, 17, 18, 19, Table 7</p><p>Quasipaa delacouri: Yan et al. 2021: 1–7.</p><p>Quasipaa cf. verrucospinosa 2: Suwannapoom et al. 2021: 1–12.</p><p>Material examined.</p><p>Holotype. • IEBR A.5174, adult male, collected by T. Q. Nguyen and C. T. Pham, on 11 March 2015, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=106.63831&amp;materialsCitation.latitude=16.941017" title="Search Plazi for locations around (long 106.63831/lat 16.941017)">Dong Chau-Khe Nuoc Trong Nature Reserve</a> (16°56.461'N, 106°38.299'E, at an elevation of 447 m a. s. l.), Le Thuy District, Quang Binh Province, Vietnam . Paratypes. (n = 8) • IEBR A.5181, adult female, the same collection data as for holotype; IEBR A. 5178, 5179, two adult males and IEBR A.5175, 5180, two adult females, collected by T. Q. Nguyen and C. T. Pham, on 18 March 2015, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=106.60947&amp;materialsCitation.latitude=16.987883" title="Search Plazi for locations around (long 106.60947/lat 16.987883)">Dong Chau - Khe Nuoc Trong Nature Reserve</a> (16°59.273'N, 106°36.568'E, at an elevation of 382 m a. s. l.), Le Thuy District, Quang Binh Province, Vietnam ; • IEBR A.5182, adult male, collected by C. T. Pham and C. V. Hoang, on 27 July 2015, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=106.62507&amp;materialsCitation.latitude=16.9506" title="Search Plazi for locations around (long 106.62507/lat 16.9506)">Dong Chau-Khe Nuoc Trong Nature Reserve</a> (16°57.036'N, 106°37.504'E, at an elevation of 300 m a. s. l.), Le Thuy District, Quang Binh Province, Vietnam ; • IEBR A.5183, adult male and IEBR A.5184, adult female, collected by C. T. Pham and T. V. Nguyen, on 5 June 2017, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=107.48437&amp;materialsCitation.latitude=16.071766" title="Search Plazi for locations around (long 107.48437/lat 16.071766)">Sao La Nature Reserve</a> (16°04.306'N, 107°29.062'E, at an elevation of 750 m a. s. l.), A Luoi District, Thua Thien Hue Province, Vietnam .</p><p>Diagnosis.</p><p>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.</p><p>Description of holotype.</p><p>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.</p><p>Forelimbs: arms short; upper arm length (UAL) 17.2 mm, forearm length (FAL) 45.1 mm; relative finger lengths: II &lt;I &lt;IV &lt;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.</p><p>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 &lt;II &lt;V &lt;III &lt;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.</p><p>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.</p><p>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.</p><p>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.</p><p>Coloration in preservative: coloration in preservative is the same in life but somewhat faded.</p><p>Sexual dimorphism.</p><p>Measurements and morphological characters of the type series are provided in Table 7. The male specimens have spines on dorsum, upper flanks, upper lip, and dorsal surface of fore- and hindlimbs small, scattered. The females contained yellowish cream eggs with melanic poles.</p><p>Ecological notes.</p><p>Specimens were found between 19: 00 and 23: 00 in the headwaters of rocky streams (Fig. 20 A). They were found in the water or on the ground of stream banks at elevations between 300 and 750 m a. s. l. The surrounding habitat was secondary forest of large, medium-sized, and small hardwoods mixed with shrubs and vines (Fig. 20 B). Air temperatures at the sites ranged from 20.1–25.7 ° C and relative humidity was 83–95 %. Male advertisement calls and tadpoles of the species had not been recorded during our field surveys. Other amphibian species found at the sites included Leptobrachium chapaense (Bourret, 1937), Xenophrys truongsonensis Luong, Hoang, Pham, Nguyen, Orlov, Ziegler &amp; Nguyen, 2022, Limnonectes kiziriani Pham, Le, Ngo, Ziegler &amp; Nguyen, 2018, L. poilani (Bourret, 1942), Amolops compotrix (Bain, Stuart &amp; Orlov, 2006), Papurana attigua (Inger, Orlov &amp; Darevsky, 1999); Odorrana gigatympana (Orlov, Ananjeva &amp; Ho, 2006), O. khalam (Stuart, Orlov &amp; Chan-ard, 2005), Hylarana maosonensis Bourret, 1937, and Rhacophorus orlovi Ziegler &amp; Köhler, 2001 .</p><p>Distribution.</p><p>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).</p><p>Comparisons.</p><p>We compared the new species with its congeners. Quasipaa binhi sp. nov. differs from Q. ohlerae sp. nov. by having the dorsum with thin and elongate ridges (vs with thick ridges); the absence nuptial spines on all fingers and ventral surface of forelimbs in males (vs present); the absence spines on lower flanks, ventral surface of forelimbs, lower lip, throat, chest, 2 / 3 anterior part of belly in males (vs present); and females with melanic pole eggs (vs wholly unpigmented); and a smaller ratio of TYE / TD in males (1.22, n = 5 vs 1.68, n = 7).</p><p>Quasipaa binhi sp. nov. differs from Q. verrucospinosa by the absence nuptial spines on all fingers in males (vs present of nuptial spines on fingers I, II, III); dorsum with thin and elongate ridges (vs dorsum with thick ridges); a smaller ratio of TD / ED (0.37, n = 5 in males and 0.38, n = 4 in females vs 0.53, n = 7 in males and 0.50, n = 7 in females); a greater ratio of TYE / TD (1.22, n = 5 in males and 1.64, n = 64 in females vs 1.16, n = 7 in males and 1.14, n = 7 in females); inner metatarsal tubercle oval (vs inner metatarsal tubercle round); different dorsal color pattern (dark brown vs yellowish grey); different ventral color pattern (immaculate white vs pale yellow); and iris dark green (vs pale copper).</p><p>Quasipaa binhi sp. nov. differs from Q. acanthophora by having the dorsum with thin and elongate ridges (vs small tubercles); the absence nuptial spines on all fingers in males (vs present on fingers I, II, III); the absence of spines on throat and chest in males (vs present); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa binhi sp. nov. differs from Q. boulengeri by the absence nuptial spines on all fingers in males (vs present on fingers I, II, III); the absence of spines on chest and belly in males (vs present); different ventral color pattern (immaculate white vs pale yellow); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa bìnhi sp. nov. differs from Q. courtoisi by having a smaller size in males (SVL 76.9–101.1 mm, n = 5 vs 126 mm, n = 1); the absence of spines on chest in males (vs present); and the absence nuptial spines on all fingers in males (vs present of nuptial spines on fingers I, II, III).</p><p>Quasipaa binhi sp. nov. differs from Q. delacouri by having the dorsum with thin and elongate ridges (vs smooth); dorsal surface of forelimbs and hindlimbs with thin and elongate ridges intermixed with small tubercles (vs smooth); different dorsal color pattern (dark brown vs brick red with black spots); and dorsum, upper flanks, upper lip, and dorsal surface of fore- and hindlimbs with small spines, scattered in males (vs absent); a greater ratio of TD / ED (0.37, n = 4 in males and 0.38, n = 4 in females vs 0.26, n = 3 in males and 0.24, n = 3 in females); a smaller ratio of TYE / TD (1.22, n = 5 in males and 1.64, n = 4 in females vs 2.15, n = 3 in males and 1.93, n = 3 in females) (Figs 13 – 15, Table 5).</p><p>Quasipaa binhi sp. nov. differs from Q. exilispinosa by having a larger size in males (SVL 76.9–101.1 mm, n = 5 in males and 88.5–123.4 mm, n = 4 in females vs SVL 44.2–66.5 mm, n = 20 in males and 40.0– 63.3 mm, n = 20 in females); the dorsum with thin and elongate ridges (vs small tubercles); the absence of spines on chest of males (vs present); the absence nuptial spines on all fingers in males (vs present on fingers I, II, III); different ventral color pattern (immaculate white vs pale yellow); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa binhi sp. nov. differs from Q. fasciculispina by having a smaller ratio of TYE / TD in males (1.22, n = 5 vs 2.0, n = 1); the absence of spines on chest in males (vs each chest tubercle with 5–10 black spines); the absence nuptial spines on all fingers in males (vs present of nuptial spines on fingers I, II, III); iris dark green (vs bright copper-colored); and external vocal sac absent (vs vocal sac openings in floor of mouth).</p><p>Quasipaa binhi sp. nov. differs from Q. jiulongensis by having the dorsum with thin and elongate ridges (vs small tubercles); the absence of spines on chest of males (vs present); the absence nuptial spines on all fingers in males (vs present on fingers I and II); the absence of pale-colored longitudinal stripes on upper jaw edge (vs present); and the absence of 4 or 5 yellow dorsal dots arranged in longitudinal rows (vs present).</p><p>Quasipaa binhi sp. nov. differs from Q. robertingeri by having different dorsal pattern (dark brown vs pale yellowish grey); the absence of spines on chest of males (vs present); the absence nuptial spines on all fingers in males (vs present of nuptial spines on fingers I, II, III); the absence of spines on chest and belly of males (vs present); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa binhi sp. nov. differs from Q. shini by having the dorsum with thin and elongate ridges (vs dorsum with thick ridges); the absence nuptial spines on all fingers in males (vs present on fingers I, II, III); and the absence of spines on chest of males (vs each chest tubercle with 3–8 black spines).</p><p>Quasipaa binhi sp. nov. differs from Q. spinosa by having the dorsum with thin and elongate ridges and round tubercles (vs small tubercles); the absence of pale-colored longitudinal stripes on upper jaw edge (vs present); the absence nuptial spines on all fingers in males (vs present on fingers I and II); and the absence of spines on chest of males (vs present).</p><p>Quasipaa binhi sp. nov. differs from Q. taoi by its larger size in females (88.5–123.4 mm, n = 4, in females vs 64.6–69.9 mm, n = 3 in females); the dorsum with thin and elongate ridges (vs dorsum with thick ridges); the absence nuptial spines on all fingers and ventral surface of forelimbs in males (vs present); and iris dark green (vs dark copper).</p><p>Quasipaa binhi sp. nov. differs from Q. yei by its larger size in males (SVL 76.8–101.1 mm, n = 5 in males and 88.5–123.4 mm, n = 4 in females vs 49.7–64.0 mm, n = 25 in males and 69.0–83.0 mm, n = 25 in females); dorsum with thin and elongate ridges (vs small tubercles); different dorsal pattern (dark brown vs pale yellowish brown); and the absence of nuptial spines around and inside vent (vs present).</p><p>Etymology.</p><p>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 ”.</p></div>	https://treatment.plazi.org/id/0C443FD8DB6F5ADB8080B11DBD3A9B6C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Pham, Cuong The;Hoang, Chung Van;Phan, Tien Quang;Pham, Anh Van;Ong, An Vinh;Nguyen, Vien Hong Thi;Ziegler, Thomas;Nguyen, Truong Quang	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.text	0A83E8F40FF55FB3A0D4CB2DDB9125C1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Quasipaa ohlerae Pham & Hoang & Phan & Pham & Ong & Nguyen & Ziegler & Nguyen 2025	<div><p>Quasipaa ohlerae sp. nov.</p><p>Figs 7, 8, 9, 10, 11, Table 6</p><p>Paa verrucospinosa: Hu et al. 2005: 340–341.</p><p>Quasipaa verrucospinosa: Yan et al. 2021: 1–7. Suwannapoom et al. 2021: 1–12, fig. 3.</p><p>Quasipaa cf. verrucospinosa: Dau et al. 2024: 9–11, fig. 5.</p><p>Type material.</p><p>Holotype. • IEBR A.5159, adult male, collected by T. Q. Phan and H. Q. Nguyen, on 17 November 2021, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.98492&amp;materialsCitation.latitude=19.9846" title="Search Plazi for locations around (long 104.98492/lat 19.9846)">Xuan Lien Nature Reserve</a> (19°59.076'N, 104°59.095'E, at an elevation of 806 m a. s. l.), Thuong Xuan District, Thanh Hoa Province, Vietnam . Paratypes. (n = 12) • IEBR A.5160, adult female, collected by T. Q. Phan and C. V. Hoang, on 20 October 2021, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Xuan Lien Nature Reserve</a> (19°52.041'N, 105°12.569'E, at an elevation of 297 m a. s. l.), Thuong Xuan District, Thanh Hoa Province, Vietnam ; IEBR A.5161 –5163, three adult males and IEBR A.5164 –5166, three adult females, collected by C. T. Pham and C. V. Hoang, on 25 August 2012, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Xuan Lien Nature Reserve</a> (19°51.446'N, 105°12.153'E, at an elevation of 423 m a. s. l.), Thuong Xuan District, Thanh Hoa Province, Vietnam ; ZVNU 11, adult male, collected by A. V. Pham, on 22 December 2012, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Copia Nature Reserve</a> (21°20.216'N, 103°34.822'E, at an elevation of 950 m a. s. l.), Thuan Chau District, Son La Province, Vietnam ; ZVNU 12, adult male, collected by T. Q. Nguyen, A. V. Pham, and H. N. Ngo, on 17 September 2014, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Copia Nature Reserve</a> (21°20.216'N, 103°34.822'E, at an elevation of 950 m a. s. l.), Thuan Chau District, Son La Province, Vietnam ; ZVNU 14, adult male and ZVNU 13, adult female, collected by A. V. Pham and N. B. Sung, on 16 September 2016, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Copia Nature Reserve</a> (21°20.216'N, 103°34.822'E, at an elevation of 950 m a. s. l.), Thuan Chau District, Son La Province, Vietnam ; IEBR A. 5167, adult female, collected by A. V. Ong, on 26 October 2021, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.9579&amp;materialsCitation.latitude=19.737417" title="Search Plazi for locations around (long 104.9579/lat 19.737417)">Pu Hoat Nature Reserve</a> (19°44.245'N, 104°57.474'E, at an elevation of 655 m a. s. l.), Que Phong District, Nghe An Province, Vietnam .</p><p>Diagnosis.</p><p>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.</p><p>Description of holotype.</p><p>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.</p><p>Forelimbs: arms short; upper arm length (UAL) 23.1 mm, forearm length (FAL) 51.0 mm; relative finger lengths: II &lt;I &lt;IV &lt;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.</p><p>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 &lt;II &lt;V &lt;III &lt;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.</p><p>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.</p><p>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.</p><p>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.</p><p>Coloration in preservative: coloration in preservative is the same in life but somewhat faded.</p><p>Sexual dimorphism.</p><p>Measurements and morphological characteristics of the type series are provided in Table 6. The male specimens have a nuptial pad on fingers I and II and black spines on lower flanks, ventral surface of forelimbs, lower lip, throat, chest, 2 / 3 anterior part of belly, and all fingers. The females contained yellowish cream eggs, wholly unpigmented.</p><p>Ecological notes.</p><p>Specimens were found between 19: 00 and 23: 00 in the headwaters of rocky streams (Fig. 12 A). They were found in the water or on the ground of stream banks at elevations between 300 and 950 m a. s. l. The surrounding habitat was secondary forest of large, medium-sized, and small hardwoods mixed with shrubs and vines (Fig. 12 B). Air temperatures at the sites ranged from 20.3–27.8 ° C and relative humidity was 65–83 %. Male advertisement calls and tadpoles of the species had not been recorded during our field surveys. Other amphibian species found at the sites included Leptobrachella eos (Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler &amp; Dubois, 2011), Xenophrys lancangica Lyu, Wang &amp; Wang, 2023, Limnonectes bannaensis Ye, Fei, Xie &amp; Jiang, 2007, Amolops tanfuilianae Sheridan, Phimmachak, Sivongxay &amp; Stuart, 2023, Odorrana chloronota (Günther, 1876), O. nasica (Boulenger, 1903), O. tiannanensis (Yang &amp; Li, 1980), Hylarana maosonensis Bourret, 1937, Kurixalus sp., Polypedates megacephalus Hallowell, 1861, and Rhacophorus orlovi Ziegler &amp; Köhler, 2001 . During to field surveys in Vu Quang National Park (Ha Tinh Province) and Pu Hoat Nature Reserve (Nghe An Province) in April 2025, We also observed some individuals of the new species.</p><p>Distribution.</p><p>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).</p><p>Comparisons.</p><p>We compared the new species with its congeners. Quasipaa ohlerae sp. nov. differs from Q. verrucospinosa by having nuptial spines on all fingers of males (vs absent on finger IV); males with nuptial spines on ventral surface of arms (vs absent); dense spines on lower lip and throat of males (vs small and scattered); a smaller ratio of TD / ED (0.32, n = 7 in males and 0.34, n = 6 in females vs 0.53, n = 7 in males and 0.50, n = 7 in females); a greater ratio of TYE / TD (1.68, n = 7 in males and 1.61, n = 6 in females vs 1.16, n = 7 in males and 1.14, n = 7 in females), inner metatarsal tubercle oval (vs inner metatarsal tubercle round); different dorsal color pattern (dark brown vs yellowish grey); different ventral color pattern (immaculate white vs pale yellow); iris dark green (vs pale copper); and females with wholly unpigmented eggs (vs melanic poles).</p><p>Quasipaa ohlerae sp. nov. differs from Q. acanthophora by having the dorsum with thick ridges (vs small tubercles); males with nuptial spines on ventral surface of arms (vs absent); males with nuptial spines on all fingers (vs absent on finger IV); dense spines present on throat and chest of males (vs small and scattered); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa ohlerae sp. nov. differs from Q. boulengeri by having the dorsum with thick ridges and round tubercles (vs elongate ridges), males with nuptial spines on all fingers (vs absent on finger IV), males with nuptial spines on throat and ventral surface of arms (vs absent), different ventral color pattern (immaculate white vs pale yellow), and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa ohlerae sp. nov. differs from Q. courtoisi by having a smaller size in males (SVL 86.7–107.8 mm, n = 7 vs 126 mm, n = 1), males with nuptial spines on throat and ventral surface of arms (vs absent) and males with nuptial spines on all fingers (vs absent on finger IV).</p><p>Quasipaa ohlerae sp. nov. differs from Q. delacouri by having the dorsum with thick ridges and round tubercles (vs smooth); males with nuptial pad on fingers I and II (vs absent); and males with nuptial spines (vs absent); different dorsal color pattern (dark brown vs brick red with black spots); and tibio-tarsal articulation reaching to nostril (vs tibio-tarsal articulation reaching to tip of snout), a greater ratio of TD / ED (0.32, n = 7 in males and 0.34, n = 6 in females vs 0.26, n = 3 in males and 0.24, n = 3 in females); a smaller ratio of TYE / TD (1.68, n = 7 in males and 1.61, n = 6 in females vs 2.15, n = 3 in males and 1.93, n = 3 in females) (Figs 13 – 15, Table 5).</p><p>Quasipaa ohlerae sp. nov. differs from Q. exilispinosa by having a larger size (SVL 86.7–107.8 mm, n = 7 in males and 92.7–107.0 mm, n = 6 in females vs SVL 44.2–66.5 mm, n = 20 in males and 40.0– 63.3 mm, n = 20 in females); dorsum with thick ridges (vs small tubercles); males with nuptial spines on throat and ventral surface of arms (vs absent); males with nuptial spines on all fingers (vs absent on finger IV); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa ohlerae sp. nov. differs from Q. fasciculispina by having a smaller ratio of TYE / TD in males (1.68, n = 7 vs 2.0, n = 1); each chest tubercle with only one black spine in males (vs each chest tubercle with 5–10 black spines); iris dark green (vs bright copper-colored); and external vocal sac absent (vs vocal sac openings in floor of mouth).</p><p>Quasipaa ohlerae sp. nov. differs from Q. jiulongensis by having the dorsum with thick ridges (vs small tubercles); males with nuptial spines on throat and ventral surface of arms of males (vs absent); males with nuptial spines on all fingers (vs absent on fingers III and IV); the absence of pale-colored longitudinal stripes on upper jaw edge (vs present); and the absence of four or five yellow dorsal dots arranged in longitudinal rows (vs present).</p><p>Quasipaa ohlerae sp. nov. differs from Q. robertingeri by having the dorsum with thick ridges and round tubercles (vs elongate ridges); males with nuptial spines on all fingers (vs absent on finger IV); males with nuptial spines on throat and ventral surface of arms (vs absent); different ventral color pattern (immaculate white vs pale yellow); and iris dark green (vs copper on upper and greyish on lower).</p><p>Quasipaa ohlerae sp. nov. differs from Q. shini by the males having nuptial spines on all fingers (vs absent on finger IV), on throat and ventral surface of arms (vs absent), and each chest tubercle with only one black spine in males (vs each chest tubercle with 3–8 black spines).</p><p>Quasipaa ohlerae sp. nov. differs from Q. spinosa by having the dorsum with thick ridges (vs small tubercles); the absence of pale-colored longitudinal stripes on upper jaw edge (vs present); males with nuptial spines on throat and ventral surface of arms of males (vs absent); and males with nuptial spines on all fingers (vs absent on fingers III and IV).</p><p>Quasipaa ohlerae sp. nov. differs from Q. taoi by its lager size (SVL 86.7–107.8 mm, n = 7 in males and 92.7–107.0 mm, n = 6 in females vs 79.6–84.3 mm, n = 3 in males and 64.6–69.9 mm, n = 3 in females); a greater ratio of TYE / TD (1.68, n = 7 in males and 1.61, n = 6 in females vs 1.11, n = 3 in males and 1.20, n = 3 in females); the presence of nuptial spines on chest and belly in males (vs absent); iris dark green (vs dark copper); and tibio-tarsal articulation reaching to nostril (vs tibio-tarsal articulation reaching to tip of snout).</p><p>Quasipaa ohlerae sp. nov. differs from Q. yei by its larger size (SVL 86.7–107.8 mm, n = 7 in males and 92.7–107.0 mm, n = 6 in females vs 49.7–64.0 mm, n = 25 in males and 69.0–83.0 mm, n = 25 in females); males with nuptial spines on lower flanks, ventral surface of forelimbs, lower lip, throat, chest, 2 / 3 anterior part of belly (vs absent); males with nuptial spines on all fingers (vs absent); and the absence of nuptial spines around vent (vs present).</p><p>Etymology.</p><p>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 ”.</p></div>	https://treatment.plazi.org/id/0A83E8F40FF55FB3A0D4CB2DDB9125C1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Pham, Cuong The;Hoang, Chung Van;Phan, Tien Quang;Pham, Anh Van;Ong, An Vinh;Nguyen, Vien Hong Thi;Ziegler, Thomas;Nguyen, Truong Quang	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.text	C9F126A2CD8257E388FF4292FE359B20.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Quasipaa verrucospinosa (Bourret 1937)	<div><p>Quasipaa verrucospinosa (Bourret, 1937)</p><p>Figs 4, 5, 6, Table 5</p><p>Rana spinosa verrucospinosa Bourret, 1937: 8., fig. 7.</p><p>Rana verrucospinosa: Bourret 1942: 295–296, fig. 83.</p><p>Paa verrucospinosa: Inger et al. 1999: 22–23.</p><p>Quasipaa verrucospinosa: Ohler and Dubois 2006: 781.</p><p>Quasipaa cf. verrucospinosa 1: Suwannapoom et al. 2021: 1–12.</p><p>Material examined.</p><p>(n = 14) • IEBR A. 5153 –5155, three adult males and IEBR A. 5021, 5156, two adult females, collected by C. T. Pham, on 18 July 2022, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.6479&amp;materialsCitation.latitude=21.45845" title="Search Plazi for locations around (long 105.6479/lat 21.45845)">Tam Dao National Park</a> (21°27.507'N, 105°38.874'E, at an elevation of 985 m a. s. l.), Vinh Phuc Province, Vietnam ; • IEBR A. 5023, 5024, two adult males and IEBR A. 5022, adult female, collected by C. V. Hoang, on 23 November 2015, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.127464&amp;materialsCitation.latitude=22.832933" title="Search Plazi for locations around (long 105.127464/lat 22.832933)">Bac Me Nature Reserve</a> (22°49.976'N, 105°07.648'E, at an elevation of 780 m a. s. l.), Ha Giang Province, Vietnam ; • IEBR A. 5028, adult male and IEBR A. 5026, 5027, two adult females, collected by C. T. Pham and T. Q. Phan, on 25 August 2017, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.33532&amp;materialsCitation.latitude=22.571466" title="Search Plazi for locations around (long 105.33532/lat 22.571466)">Sinh Long Commune</a> (22°34.288'N, 105°20.119'E, at an elevation of 801 m a. s. l.), Na Hang District, Tuyen Quang Province ; • IEBR A. 5025, adult female, collected by C. T. Pham, C. V. Hoang, and T. Q. Phan, on 27 October 2018, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=105.073715&amp;materialsCitation.latitude=22.208233" title="Search Plazi for locations around (long 105.073715/lat 22.208233)">Cham Chu Nature Reserve</a> (22°12.494'N, 105°04.423'E; at an elevation of 981 m a. s. l.), Ham Yen District, Tuyen Quang Province ; • IEBR A. 5157, adult male and IEBR A. 5158, adult female, collected by C. T. Pham, on 12 October 2022, in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=104.0699&amp;materialsCitation.latitude=22.159283" title="Search Plazi for locations around (long 104.0699/lat 22.159283)">Hoang Lien National Park</a> (22°09.557'N, 104°04.194'E; at an elevation 2,078 m a. s. l.), Sa Pa District, Lao Cai Province, Vietnam .</p><p>Revised diagnosis.</p><p>(1) A large frog (SVL up to 106 mm in males and 95 mm in females); (2) head broader than long (HL / HW 0.88 in males, 0.89 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) males with dense spines on chest, 2 / 3 anterior part of belly and fingers I, II, III (sometimes present on fingers I and II only); (11) male without spines on finger IV and ventral surface of forelimbs; (12) finger I with nuptial pad in males; (13) yellowish cream eggs with melanic poles in females; (14) toes fully webbed to distal end of terminal phalanx; and (15) in life, dorsum yellowish grey or pale brownish grey, chest and belly pale yellowish white, iris pale copper.</p><p>Description.</p><p>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.</p><p>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 &lt;I &lt;IV &lt;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.</p><p>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 &lt;II &lt;V &lt;III &lt;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.</p><p>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.</p><p>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.</p><p>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.</p><p>Sexual dimorphism.</p><p>Measurements and morphological characters of the Quasipaa verrucospinosa are provided in Table 5. males are slightly larger than females (SVL 100.7 ± 6.6 mm, n = 7 males vs 86.8 ± 7.4 mm, n = 7 females). The male specimens have a nuptial pad on finger I and dense spines on lower part of flanks, chest, 2 / 3 anterior part of belly, and fingers I, II, III. The females contained yellowish cream eggs with melanic poles.</p><p>Distribution.</p><p>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.</p><p>Remarks.</p><p>The specimens agreed well with the descriptions of Bourret (1942) and Inger et al. (1999) in size, skin texture, and coloration; males with dense spines on 2 / 3 anterior part of belly. In addition, dorsum yellowish grey and belly pale yellowish white (more yellow in females); males with spines on fingers I, II, and III (some times present on fingers I and II only); yellowish cream eggs with melanic poles in females; iris pale copper.</p></div>	https://treatment.plazi.org/id/C9F126A2CD8257E388FF4292FE359B20	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Pham, Cuong The;Hoang, Chung Van;Phan, Tien Quang;Pham, Anh Van;Ong, An Vinh;Nguyen, Vien Hong Thi;Ziegler, Thomas;Nguyen, Truong Quang	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
