Research Article |
Corresponding author: Cuong The Pham ( cuongiebr@gmail.com ) Corresponding author: Thomas Ziegler ( ziegler@koelnerzoo.de ) Academic editor: Annemarie Ohler
© 2022 Cuong The Pham, Chung Van Hoang, Tien Quang Phan, Truong Quang Nguyen, Thomas Ziegler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pham CT, Hoang CV, Phan TQ, Nguyen TQ, Ziegler T (2022) Hidden in the jungle of Vietnam: a new species of Quasipaa (Amphibia, Anura, Dicroglossidae) from Ngoc Linh Mountain. ZooKeys 1124: 23-42. https://doi.org/10.3897/zookeys.1124.89282
|
A new species of Quasipaa is described from Ngoc Linh Mountain of the Kon Tum Massif in central Vietnam. The new species is morphologically distinguishable from its congeners on the basis of a combination of the following diagnostic characters: SVL 79.6–84.3 mm in males and 64.6–69.9 mm in females; head broader than long; vomerine teeth present; external vocal sacs absent; tympanum slightly visible; dorsum with lines of thick ridges and small round tubercles; flanks covered by oval and round tubercles; supratympanic fold present; dorsolateral fold absent; ventrolateral sides, ventral surface of arms, and all fingers with spines in males; the absence of spines on chest and belly in males; toes fully webbed to distal portion of terminal phalanx; in life, dorsum dark brown, chest and belly immaculate white. Phylogenetic analyses found that the genetic divergence of the new species and its congeners ranged from 4.2–5.1% (compared with Quasipaa boulengeri) to 7.6–8.1% (compared with Q. shini) in the 16S gene.
Kon Tum Province, molecular phylogeny, Quasipaa taoi sp. nov., taxonomy
The genus Quasipaa Dubois, 1992 is known from China through the Indochina region and south to Thailand. The genus currently contains 12 recognized species, including Quasipaa acanthophora Dubois & Ohler, 2009, Q. boulengeri (Günther, 1889), Q. courtoisi (Angel, 1922), Q. delacouri (Angel, 1928), Q. exilispinosa (Liu & Hu, 1975), Q. fasciculispina (Inger, 1970), Q. jiulongensis (Huang & Liu, 1985), Q. robertingeri (Wu & Zhao, 1995), Q. shini (Ahl, 1930), Q. spinosa (David, 1875), Q. verrucospinosa (Bourret, 1937), and Q. yei (Chen, Qu & Jiang, 2002) (
During our recent fieldwork in the Central Highlands of Vietnam, specimens of Quasipaa were collected in the evergreen forests of Ngoc Linh Mountain, Kon Tum Province. These specimens were identified as members of the “Quasipaa sensu stricto” species group (Group II-2) (
A field survey was conducted in March 2019 in Ngoc Linh Nature Reserve, Dak Giei District, Kon Tum Province. Frogs were collected between 19:00 and 23:00. After taking photographs of living specimens, they were anaesthetized and euthanized in a closed vessel with a piece of cotton wool containing ethyl acetate (
In this study, 15 samples of five species of Quasipaa were used for molecular analysis (Table
GenBank accession numbers and associated samples that used in this study.
Species | Location | Genbank No. | References | |
---|---|---|---|---|
1 | Quasipaa taoi sp. nov. | Kon Tum, Vietnam | OP326684 | This study |
2 | Quasipaa taoi sp. nov. | Kon Tum, Vietnam | OP326685 | This study |
3 | Quasipaa taoi sp. nov. | Kon Tum, Vietnam | EU979804 |
|
4 | Quasipaa taoi sp. nov. | Xekong, Laos | EU979803 |
|
5 | Q. verrucospinosa | Vinh Phuc, Vietnam | EU979813 |
|
6 | Q. verrucospinosa | Tuyen Quang, Vietnam | OP326686 | This study |
7 | Q. verrucospinosa | Tuyen Quang, Vietnam | OP326687 | This study |
8 | Q. verrucospinosa | Tuyen Quang, Vietnam | OP326688 | This study |
9 | Q. spinosa | Yunnan, China | DQ118480 |
|
10 | Q. robertingeri | Sichuan, China | EU979814 |
|
11 | Q. robertingeri | Sichuan, China | DQ118478 |
|
12 | Q. boulengeri | Cao Bang, Vietnam | OP326689 | This study |
13 | Q. boulengeri | Cao Bang, Vietnam | OP326690 | This study |
14 | Q. boulengeri | Cao Bang, Vietnam | OP326691 | This study |
15 | Q. boulengeri | Cao Bang, Vietnam | OP326692 | This study |
16 | Q. boulengeri | Cao Bang, Vietnam | OP326693 | This study |
17 | Q. exilispinosa | Fujian, China | DQ118484 |
|
18 | Q. jiulongensis | Fujian, China | KF199149 |
|
19 | Q. acanthophora | Lang Son, Vietnam | OP326694 | This study |
20 | Q. acanthophora | Lang Son, Vietnam | OP326695 | This study |
21 | Q. yei | Henan, China | DQ118488 |
|
22 | Q. shini | Guangxi, China | DQ118487 |
|
23 | Quasipaa sp. | Xekong, Laos | EU979812 |
|
24 | Q. delacouri | Tuyen Quang, Vietnam | OP326696 | This study |
25 | Q. delacouri | Tuyen Quang, Vietnam | OP326697 | This study |
26 | Q. delacouri | Tuyen Quang, Vietnam | OP326698 | This study |
Outgroup | ||||
Fejervarya limnocharis | Vinh Phuc, Vietnam | EU979847 |
|
In addition, we used 11 available sequences of 16S rRNA of nine species of the genus Quasipaa in GenBank for phylogenetic analyses (
Phylogenetic trees were constructed by using maximum likelihood (ML) and Bayesian inference (BI). Chromas Pro software (Technelysium Pty Ltd., Tewantin, Australia) was used to edit the sequences, which were aligned using the ClustalW (
Measurements were taken with digital calipers to the nearest 0.1 mm. The following abbreviations were used:
SVL snout–vent length;
HL head length (measured as a parallel line with the vertebral column from posterior margin of mandible to tip of snout);
HW maximum head width (at rictus);
RL rostral length (from anterior corner of orbit to tip of snout);
NS distance from nostril to tip of snout;
EN distance from anterior corner of orbit to nostril;
IND internarial distance;
IOD interorbital distance;
ED eye diameter;
UEW maximum width of upper eyelid;
DAE distance between anterior margins of orbits;
DPE distance between posterior margins of orbits;
MN posterior margin of mandible to nostril;
MFE posterior margin of mandible to anterior margin of orbit;
MBE posterior margin of mandible to posterior margin of eye;
TD tympanum diameter;
TYE distance from anterior margin of tympanum to posterior corner of orbit;
UAL upper arm length (from axilla to elbow);
FAL forearm length (from elbow to tip of third finger);
FL1–4 finger length I–IV (from inner to outer);
NPL nuptial pad length - finger I;
FeL femur length (from vent to knee);
TbL tibia length (from knee to tarsus);
TbW maximum tibia width;
FoL foot length (from tarsus to tip of fourth toe);
TL 1–5 toe length I–V;
IMT inner metatarsal tubercle length.
For webbing formula, we followed
Morphological data were obtained by comparison of the new species with specimens of other members of the genus Quasipaa (see Appendix
Measurements were used to compare the morphometric difference between the new species from Kon Tum Province (three males and three females) vs Quasipaa boulengeri from Cao Bang Province (six males and five females). All statistical analyses were performed using PAST v. 2.17b software (
The combined matrix contained 495 aligned characters. Of those, 416 sites were conserved, and 78 sites were variable, of which 62 were found to be potentially parsimony informative. The estimated Transition/Transversion bias (R) is 3.86. Substitution pattern and rates were estimated under the
Uncorrected p-distance matrix showing percentage pairwise genetic divergences (%) for the 16SrRNA gene between members of the genus Quasipaa.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | Quasipaa taoi sp. nov. | 0.0–0.8 | |||||||||||
2 | Q. verrucospinosa | 6.4–7.1 | 0.0–0.2 | ||||||||||
3 | Q. spinosa | 6.7–7.1 | 3.1–3.3 | 0.0 | |||||||||
4 | Q. robertingeri | 5.3–5.8 | 5.3–5.5 | 4.4 | 0.0 | ||||||||
5 | Q. boulengeri | 4.2–5.1 | 5.1–5.8 | 4.6–4.9 | 1.6–1.9 | 0.0–0.4 | |||||||
6 | Q. exilispinosa | 6.7–6.9 | 6.5–6.7 | 6.7 | 6.2 | 4.9–5.1 | 0.0 | ||||||
7 | Q. jiulongensis | 4.9–5.5 | 5.5–5.8 | 5.8 | 4.7 | 3.8–4.0 | 3.3 | 0.0 | |||||
8 | Q. acanthophora | 6.2–6.9 | 7.1–7.6 | 7.8–8.0 | 7.1–7.4 | 5.8–6.2 | 2.7–2.9 | 3.3–3.5 | 0.0–0.2 | ||||
9 | Q. yei | 5.8–6.4 | 6.0–6.2 | 6.4 | 5.1 | 4.7–4.9 | 5.8 | 5.1 | 5.3–5.5 | 0.0 | |||
10 | Q. shini | 7.6–8.1 | 7.4–7.6 | 7.8 | 6.4 | 6.2–6.7 | 7.2 | 6.0 | 7.0–7.2 | 5.6 | 0.0 | ||
11 | Quasipaa sp. | 4.9–5.6 | 6.0–6.2 | 6.2 | 4.7 | 3.6–4.0 | 5.1 | 3.4 | 4.2–4.4 | 4.5 | 5.6 | 0.0 | |
12 | Q. delacouri | 4.9–5.8 | 5.8–6.0 | 6.7 | 5.1 | 4.0–4.5 | 5.1 | 4.2 | 4.2–4.5 | 3.8 | 5.6 | 1.4 | 0.0 |
The ML and BI analyses produced topologies with –ln L = 1672.0337 and 1729.0216, respectively, with a gamma shape parameter (G: 0.1363 in ML and 0.1767 in BI). Phylogenetic analyses employing ML and BI methods were nearly identical, with most well-supported nodes on the ML tree also well-supported on the BI tree, and only the BI tree is presented in Fig.
Our phylogenetic results were in general agreement with those supported by analyses in
In the following, based on the distinct molecular divergence in concert with diagnostic morphological differences compared to congeners, we describe the Quasipaa population from Ngoc Linh based on our integrative taxonomic analysis, as new species to science.
IEBR A.4997, adult male, collected by T. Q. Phan and T. D. Tran on 6 March 2019 (15°05'23.3"N, 107°51'17.5"E, at an elevation of 1,560 m asl.) in the evergreen forest of Ngoc Linh Natural Reserve, Xop Commune, Dak Glei District, Kon Tum Province, Vietnam.
IEBR A.4998, adult male; IEBR A.4999, adult male; IEBR A.5000, adult female; IEBR A.5037, adult female; IEBR A.5038, adult female, the same data as the holotype.
Both morphological characters (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 (see
A large frog (SVL 84.3 mm); habitus robust with enlarged head (HL/SVL 0.40, HW/SVL 0.43); head broader than long (HL 33.5 mm, HW 36.3 mm); snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to eye than to the tip of snout (NS 7.6 mm, EN 5.5 mm); canthus rostralis indistinct; loreal region oblique and slightly concave; rostral length greater than eye diameter (RL 13.1 mm, ED 9.7 mm); internarial distance wider than interorbital distance and upper eyelid width (IND 8.6 mm, IOD 6.2 mm, UEW 7.7 mm); tympanum slightly visible (TYD 4.1 mm) smaller than the distance from tympanum to eye (TYE 4.9 mm); vomerine teeth in two oblique ridges; tongue cordiform, notched posteriorly; external vocal sac absent.
Forelimbs: arms short; upper arm length (UAL) 17.1 mm, forearm length (FAL) 41.5 mm; relative finger lengths: II<I<IV<III; fingers free of webbing; dermal ridge on sides of fingers present on fingers I, II, III; 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.
Hindlimbs: tibia length longer than thigh length (FeL 44.2 mm, TbL 49.7 mm), approximately 3.4 times longer than wide (TbW 14.5 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 tip of snout.
Skin texture in life: dorsal surface of head with oval and round tubercles, dorsum with six lines of 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 chest, belly, and ventral surface of hindlimbs; dense spines on dorsum, flanks, ventral surface of forelimbs, ventrolateral sides, and fingers I, II; spines present on throat, dorsal surface of fore- and hindlimbs, and fingers III, IV, small and scattered.
Coloration in life: iris dark copper; dorsum and upper part of flanks dark brown; lower part of flanks whitish brown with white tubercles and black spines on top; supratympanic fold dark brown; dorsal surface of limbs yellowish brown with dark crossbars; ventral surface of limbs light yellow with brown markings; throat white with brown markings; chest and belly immaculate white; toe webbing dark brown.
Coloration in preservative: coloration in preservative is the same in life but somewhat faded.
Measurements and morphological characters of the type series are provided in Table
Measurements (in mm) and proportions of the type series of Quasipaa boulengeri and Quasipaa taoi sp. nov. (H = holotype, P = paratype, SD = standard deviation; for other abbreviations see Material and methods).
Quasipaa taoi sp. nov. | ||||||||||
Specimen ID | IEBR A.4997 | IEBR A.4999 | IEBR A.4998 | Min–Max (n = 3) | Mean±SD (n = 3) | IEBR A.5000 | IEBR A.5038 | IEBR A.5037 | Min–Max (n = 3) | Mean±SD (n = 3) |
Sex | ♂ | ♂ | ♂ | ♀ | ♀ | ♀ | ||||
Type status | H | P | P | P | P | P | ||||
SVL | 84.3 | 79.6 | 84.2 | 79.6–84.3 | 82.7±2.69 | 68.2 | 69.9 | 64.6 | 64.6–69.9 | 67.6±2.7 |
HL | 33.5 | 31.3 | 32.2 | 31.3–33.5 | 32.33±1.11 | 27.1 | 28.4 | 24.9 | 24.9–28.4 | 26.8±1.7 |
HW | 36.3 | 34.9 | 36.7 | 34.9–36.7 | 35.97±0.95 | 28.3 | 31.2 | 27.8 | 27.8–31.2 | 29.1±1.8 |
MN | 27.5 | 26.1 | 25.9 | 25.9–27.5 | 26.5±0.87 | 22.5 | 25.0 | 21.3 | 21.3–25 | 22.9±1.9 |
MFE | 22.9 | 21.9 | 22.2 | 21.9–22.9 | 22.33±0.51 | 18.8 | 20.1 | 17.6 | 17.6–20.1 | 18.8±1.3 |
MBE | 13.8 | 13.7 | 12.3 | 12.3–13.8 | 13.27±0.84 | 12.1 | 12.0 | 9.9 | 9.9–12.1 | 11.3±1.2 |
RL | 13.1 | 12.8 | 13.4 | 12.8–13.4 | 13.1±0.30 | 11.1 | 11.5 | 9.7 | 9.7–11.5 | 10.8±0.9 |
ED | 9.7 | 9.6 | 10.9 | 9.6–10.9 | 10.07±0.72 | 8.5 | 7.9 | 7.7 | 7.7–8.5 | 8±0.4 |
UEW | 7.7 | 7.1 | 7.6 | 7.1–7.7 | 7.47±0.32 | 6.2 | 7.4 | 6.0 | 6–7.4 | 6.5±0.8 |
IND | 8.6 | 7.8 | 8.5 | 7.8–8.6 | 8.3±0.44 | 7.1 | 6.2 | 5.1 | 5.1–7.1 | 6.1±1 |
IOD | 6.2 | 6.1 | 5.8 | 5.8–6.2 | 6.03±0.21 | 4.8 | 4.9 | 4.8 | 4.8–4.9 | 4.8±0.1 |
DAE | 12.2 | 13.2 | 13.6 | 12.2–13.6 | 13.0±0.72 | 11.3 | 11.1 | 9.0 | 9–11.3 | 10.5±1.3 |
DPE | 23.7 | 22.1 | 23.5 | 22.1–23.7 | 23.1±0.87 | 18.5 | 20.4 | 17.8 | 17.8–20.4 | 18.9±1.3 |
NS | 7.6 | 7.5 | 7.9 | 7.5–7.9 | 7.67±0.21 | 6.5 | 7.2 | 6.5 | 6.5–7.2 | 6.7±0.4 |
EN | 5.5 | 5.3 | 5.4 | 5.3–5.5 | 5.4±0.10 | 4.6 | 4.9 | 4.1 | 4.1–4.9 | 4.5±0.4 |
TD | 4.1 | 4.4 | 4.8 | 4.1–4.8 | 4.43±0.35 | 3.4 | 4.6 | 3.8 | 3.4–4.6 | 3.9±0.6 |
TYE | 4.9 | 5 | 4.9 | 4.9–5.0 | 4.93±0.06 | 4.7 | 5.0 | 4.3 | 4.3–5 | 4.7±0.4 |
UAL | 17.1 | 17.6 | 19.2 | 17.1–19.2 | 17.97±1.10 | 13.6 | 11.7 | 11.1 | 11.1–13.6 | 12.1±1.3 |
FAL | 41.5 | 39.9 | 41.2 | 39.9–41.5 | 40.87±0.85 | 33.2 | 33.5 | 29.6 | 29.6–33.5 | 32.1±2.1 |
FeL | 44.2 | 45.3 | 44.6 | 44.2–45.3 | 44.7±0.56 | 33.8 | 36.5 | 33.6 | 33.6–36.5 | 34.6±1.6 |
TbL | 49.7 | 48.9 | 48.3 | 48.3–49.7 | 48.97±0.7 | 40.1 | 41.0 | 37.7 | 37.7–41 | 39.6±1.7 |
TbW | 14.5 | 15.8 | 15.5 | 14.5–15.8 | 15.27±0.68 | 10.1 | 12.4 | 11.4 | 10.1–12.4 | 11.3±1.1 |
FoL | 65.6 | 63.3 | 64.9 | 63.3–65.6 | 64.6±1.18 | 55.7 | 54.6 | 50.1 | 50.1–55.7 | 53.5±3 |
IMT | 6.3 | 5.9 | 6.1 | 5.9–6.3 | 6.1±0.20 | 4.8 | 4.6 | 4.6 | 4.6–4.8 | 4.7±0.1 |
HL/SVL | 0.4 | 0.39 | 0.38 | 0.38–0.40 | 0.39±0.01 | 0.39 | 0.39 | 0.41 | 0.39–0.41 | 0.4±0.01 |
HW/SVL | 0.43 | 0.44 | 0.44 | 0.43–0.44 | 0.43±0.00 | 0.43 | 0.41 | 0.45 | 0.41–0.45 | 0.43±0.02 |
RL/SVL | 0.16 | 0.16 | 0.16 | 0.16–0.16 | 0.16±0.0 | 0.15 | 0.15 | 0.16 | 0.15–0.16 | 0.16±0.01 |
HL/HW | 0.92 | 0.9 | 0.88 | 0.88–0.92 | 0.9±0.02 | 0.90 | 0.90 | 0.96 | 0.9–0.96 | 0.92±0.03 |
ED/RL | 0.74 | 0.75 | 0.81 | 0.74–0.81 | 0.77±0.04 | 0.79 | 0.69 | 0.79 | 0.69–0.79 | 0.75±0.06 |
TYE/TD | 1.2 | 1.14 | 1.02 | 1.02–1.20 | 1.11±0.16 | 1.13 | 1.09 | 1.38 | 1.09–1.38 | 1.2±0.16 |
ED/TD | 0.42 | 0.46 | 0.44 | 0.42–0.46 | 0.44±0.02 | 0.49 | 0.4 | 0.58 | 0.4–0.58 | 0.49±0.01 |
TbL/SVL | 0.59 | 0.61 | 0.57 | 0.57–0.61 | 0.59±0.02 | 0.58 | 0.58 | 0.59 | 0.58–0.59 | 0.59±0 |
TbL/TbW | 3.43 | 3.09 | 3.12 | 3.09–3.43 | 3.21±0.19 | 3.30 | 3.30 | 3.97 | 3.3–3.97 | 3.53±0.38 |
Q. boulengeri | ||||||||||
Specimen ID | IEBR A.5007 | IEBR A.5008 | IEBR A.5009 | IEBR A.50109 | IEBR A.5011 | IEBR A.5012 | Min–Max (n = 6) | Mean±SD (n = 6) | ||
Sex | ♂ | ♂ | ♂ | ♂ | ♂ | ♂ | ||||
Type status | ||||||||||
SVL | 100.7 | 96.8 | 99.2 | 101.7 | 87.8 | 92.6 | 87.8–101.7 | 96.5±5.3 | ||
HL | 37.3 | 37.5 | 39.8 | 39.2 | 35.8 | 36.3 | 35.8–39.8 | 37.6±1.6 | ||
HW | 42.0 | 43.1 | 45.3 | 44.8 | 40.1 | 40.9 | 40.1–45.3 | 42.7±2.1 | ||
MN | 31.6 | 31.7 | 34.5 | 34.4 | 30.2 | 31.0 | 30.2–34.5 | 32.2±1.8 | ||
MFE | 26.1 | 25.9 | 27.9 | 28.8 | 24.7 | 25.8 | 24.7–28.8 | 26.5±1.5 | ||
MBE | 17.4 | 16.4 | 17.7 | 19.5 | 15.5 | 15.6 | 15.5–19.5 | 17±1.5 | ||
RL | 14.9 | 14.3 | 15.7 | 14.6 | 13.9 | 14.7 | 13.9–15.7 | 14.7±0.6 | ||
ED | 10.2 | 10.0 | 12.4 | 10.7 | 10.2 | 10.4 | 10–12.4 | 10.6±0.9 | ||
UEW | 8.3 | 7.4 | 8.0 | 7.9 | 7.3 | 6.5 | 6.5–8.3 | 7.6±0.7 | ||
IND | 8.3 | 7.9 | 9.3 | 8.2 | 8.3 | 8.0 | 7.9–9.3 | 8.3±0.5 | ||
IOD | 6.6 | 5.9 | 8.4 | 8.2 | 5.5 | 7.5 | 5.5–8.4 | 7±1.2 | ||
DAE | 14.7 | 14.3 | 15.8 | 15.6 | 13.0 | 13.5 | 13–15.8 | 14.5±1.1 | ||
DPE | 26.1 | 25.4 | 29.6 | 28.4 | 24.3 | 26.0 | 24.3–29.6 | 26.6±2 | ||
NS | 9.1 | 7.0 | 7.7 | 7.4 | 7.7 | 7.8 | 7–9.1 | 7.8±0.7 | ||
Q. boulengeri | ||||||||||
Specimen ID | IEBR A.5007 | IEBR A.5008 | IEBR A.5009 | IEBR A.50109 | IEBR A.5011 | IEBR A.5012 | Min–Max (n = 6) | Mean±SD (n = 6) | ||
EN | 6.3 | 6.7 | 8.2 | 7.3 | 5.8 | 6.5 | 5.8–8.2 | 6.8±0.8 | ||
TD | 6.5 | 6.0 | 6.7 | 6.3 | 6.1 | 5.7 | 5.7–6.7 | 6.2±0.4 | ||
TYE | 5.6 | 5.6 | 7.1 | 6.2 | 5.0 | 6.2 | 5–7.1 | 5.9±0.7 | ||
UAL | 20.0 | 19.3 | 21.0 | 20.0 | 15.4 | 15.7 | 15.4–21 | 18.6±2.4 | ||
FAL | 46.0 | 49.6 | 51.9 | 52.8 | 46.5 | 45.7 | 45.7–52.8 | 48.7±3.1 | ||
FeL | 54.1 | 50.3 | 57.0 | 52.3 | 48.2 | 48.7 | 48.2–57 | 51.8±3.4 | ||
TbL | 53.9 | 52.6 | 56.3 | 57.4 | 49.7 | 52.2 | 49.7–57.4 | 53.7±2.8 | ||
TbW | 16.8 | 17.7 | 17.7 | 19.0 | 16.5 | 17.3 | 16.5–19 | 17.5±0.9 | ||
FoL | 72.5 | 72.3 | 77.3 | 78.6 | 67.8 | 68.7 | 67.8–78.6 | 72.9±4.4 | ||
IMT | 8.1 | 8.1 | 7.4 | 8.8 | 7.7 | 8.0 | 7.4–8.8 | 8±0.5 | ||
HL/SVL | 0.37 | 0.39 | 0.40 | 0.39 | 0.41 | 0.39 | 0.37–0.41 | 0.4±0 | ||
HW/SVL | 0.42 | 0.45 | 0.46 | 0.44 | 0.46 | 0.44 | 0.42–0.46 | 0.4±0 | ||
RL/SVL | 0.15 | 0.15 | 0.16 | 0.14 | 0.16 | 0.16 | 0.14–0.16 | 0.2±0 | ||
HL/HW | 0.89 | 0.87 | 0.88 | 0.88 | 0.89 | 0.89 | 0.87–0.89 | 0.9±0 | ||
ED/RL | 0.69 | 0.70 | 0.79 | 0.73 | 0.73 | 0.71 | 0.69–0.79 | 0.7±0 | ||
TYE/TD | 0.86 | 0.93 | 1.05 | 0.98 | 0.82 | 1.09 | 0.82–1.09 | 1±0.1 | ||
ED/TD | 0.64 | 0.60 | 0.55 | 0.59 | 0.60 | 0.55 | 0.55–0.64 | 0.59±0 | ||
TbL/SVL | 0.54 | 0.54 | 0.57 | 0.56 | 0.57 | 0.56 | 0.54–0.57 | 0.6±0 | ||
TbL/TbW | 3.21 | 2.97 | 3.18 | 3.02 | 3.02 | 3.01 | 2.97–3.21 | 3.1±0.1 | ||
Q. boulengeri | ||||||||||
Specimen ID | IEBR A.5013 | IEBR A.5014 | IEBR A.5039 | IEBR A.5040 | IEBR A.5041 | Min–Max (n = 5) | Mean±SD (n = 5) | |||
Sex | ♀ | ♀ | ♀ | ♀ | ♀ | |||||
Type status | ||||||||||
SVL | 105.5 | 95.5 | 90.8 | 82.5 | 91.4 | 82.5–105.5 | 93.1±8.4 | |||
HL | 39.9 | 36.4 | 34.1 | 30.3 | 34.9 | 30.3–39.9 | 35.1±3.5 | |||
HW | 44.2 | 39.5 | 39.4 | 39.4 | 41.0 | 39.4–44.2 | 40.7±2.1 | |||
MN | 32.9 | 30.0 | 28.1 | 25.5 | 30.8 | 25.5–32.9 | 29.5±2.8 | |||
MFE | 26.7 | 24.4 | 22.1 | 19.8 | 25.0 | 19.8–26.7 | 23.6±2.7 | |||
MBE | 17.6 | 13.7 | 13.7 | 11.9 | 16.1 | 11.9–17.6 | 14.6±2.2 | |||
RL | 15.7 | 13.7 | 13.0 | 11.4 | 12.6 | 11.4–15.7 | 13.3±1.6 | |||
ED | 10.8 | 11.4 | 9.4 | 9.1 | 10.6 | 9.1–11.4 | 10.3±1 | |||
UEW | 7.7 | 7.3 | 7.6 | 6.7 | 7.9 | 6.7–7.9 | 7.4±0.5 | |||
IND | 9.4 | 7.8 | 8.0 | 7.5 | 9.0 | 7.5–9.4 | 8.3±0.8 | |||
IOD | 8.5 | 6.0 | 5.9 | 6.1 | 5.4 | 5.4–8.5 | 6.4±1.2 | |||
DAE | 15.8 | 13.5 | 12.8 | 13.0 | 12.2 | 12.2–15.8 | 13.4±1.4 | |||
DPE | 28.6 | 24.8 | 13.8 | 21.9 | 9.0 | 9–28.6 | 19.6±8 | |||
NS | 9.3 | 7.3 | 7.6 | 6.9 | 8.2 | 6.9–9.3 | 7.9±0.9 | |||
EN | 6.1 | 6.6 | 5.6 | 5.3 | 5.8 | 5.3–6.6 | 5.9±0.5 | |||
TD | 6.7 | 5.4 | 5.1 | 5.2 | 6.1 | 5.1–6.7 | 5.7±0.7 | |||
TYE | 6.6 | 4.9 | 5.0 | 3.8 | 5.7 | 3.8–6.6 | 5.2±1 | |||
UAL | 16.4 | 16.6 | 15.3 | 11.8 | 15.1 | 11.8–16.6 | 15±1.9 | |||
FAL | 47.7 | 43.0 | 40.6 | 36.7 | 43.7 | 36.7–47.7 | 42.3±4.1 | |||
FeL | 52.0 | 47.7 | 46.6 | 41.2 | 46.6 | 41.2–52 | 46.8±3.8 | |||
TbL | 54.0 | 50.5 | 47.8 | 44.2 | 49.8 | 44.2–54 | 49.3±3.6 | |||
TbW | 18.7 | 18.1 | 17.8 | 14.9 | 16.5 | 14.9–18.7 | 17.2±1.5 | |||
FoL | 74.4 | 67.6 | 65.4 | 60.0 | 69.2 | 60–74.4 | 67.3±5.3 | |||
IMT | 8.1 | 6.8 | 16.1 | 5.5 | 7.8 | 5.5–16.1 | 8.9±4.2 | |||
HL/SVL | 0.38 | 0.38 | 0.38 | 0.37 | 0.38 | 0.4–0.4 | 0.4±0 | |||
HW/SVL | 0.42 | 0.41 | 0.43 | 0.48 | 0.45 | 0.4–0.5 | 0.4±0 | |||
RL/SVL | 0.15 | 0.14 | 0.14 | 0.14 | 0.14 | 0.1–0.1 | 0.1±0 | |||
HL/HW | 0.90 | 0.92 | 0.87 | 0.77 | 0.85 | 0.8–0.9 | 0.9±0.1 | |||
ED/RL | 0.69 | 0.83 | 0.72 | 0.80 | 0.84 | 0.7–0.8 | 0.8±0.1 | |||
TYE/TD | 0.99 | 0.92 | 0.98 | 0.74 | 0.94 | 0.7–1 | 0.9±0.1 | |||
ED/TD | 0.62 | 0.47 | 0.54 | 0.57 | 0.58 | 0.47–0.62 | 0.56±0 | |||
TbL/SVL | 0.51 | 0.53 | 0.53 | 0.54 | 0.54 | 0.5–0.5 | 0.5±0 | |||
TbL/TbW | 2.88 | 2.79 | 2.69 | 2.96 | 3.02 | 2.7–3 | 2.9±0.1 |
The new species is named in honor of our colleague and friend, Assoc. Prof. Dr. Tao Thien Nguyen from the Institute of Genome Research, Vietnam Academy of Science and Technology, in recognition of his numerous scientific contributions towards a better understanding of the amphibians of Vietnam. We recommend “Tao’s Spiny Frog” as the common English name of the new species and the common name in Vietnamese as “Ếch gai sần tạo”.
Specimens were found between 19:00 and 23:00 in the headwaters of rocky streams (Fig.
Quasipaa taoi sp. nov. is currently known from Ngoc Linh Mountain of the Central Highlands in Vietnam (Fig.
We compared the new species with its congeners. Quasipaa taoi sp. nov. differs from Q. boulengeri by having a smaller size, SVL 79.6–84.3 mm, n = 3 in males, 64.6–69.9 mm, n = 3 in females (vs 87.8–101.7 mm, n = 6 in males, 82.5–105.5 mm, n = 5 in females), 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), and the absence of nuptial spines on chest and belly in males (vs present). In the PCA analysis, the first two principal component axes could separate Quasipaa taoi sp. nov. from Q. boulengeri by 24 characters (Fig.
Variable loadings for principal components with eigenvalue greater than 0.01, from morphometric characters corrected by SVL. All measurements were given in millimeter (mm).
Male | Female | |||||
---|---|---|---|---|---|---|
Axis 1 | Axis 2 | Axis 3 | Axis 1 | Axis 2 | Axis 3 | |
Eigenvalue | 0.039 | 0.004 | 0.003 | 0.118 | 0.032 | 0.012 |
% variance | 76.63 | 8.86 | 5.75 | 67.42 | 18.57 | 7.00 |
SVL | 0.189 | −0.066 | 0.117 | 0.223 | 0.055 | 0.012 |
HW | 0.205 | −0.077 | 0.039 | 0.216 | 0.033 | 0.041 |
HL | 0.181 | −0.068 | 0.005 | 0.202 | 0.056 | 0.047 |
MN | 0.234 | −0.083 | −0.055 | 0.189 | 0.033 | 0.123 |
MFE | 0.212 | −0.056 | −0.058 | 0.178 | 0.038 | 0.157 |
MBE | 0.300 | −0.154 | −0.016 | 0.212 | 0.017 | 0.224 |
RL | 0.139 | 0.009 | 0.021 | 0.173 | 0.100 | 0.005 |
ED | 0.106 | 0.258 | 0.065 | 0.164 | 0.052 | 0.117 |
UEW | 0.056 | 0.148 | 0.460 | 0.111 | −0.035 | 0.094 |
IND | 0.039 | 0.216 | 0.112 | 0.229 | −0.002 | 0.166 |
IOD | 0.261 | 0.479 | −0.442 | 0.195 | 0.207 | −0.098 |
DAE | 0.167 | 0.138 | 0.169 | 0.182 | 0.150 | −0.012 |
DPE | 0.198 | 0.109 | −0.036 | −0.008 | 0.825 | −0.441 |
NS | 0.008 | 0.013 | 0.227 | 0.134 | 0.027 | 0.108 |
EN | 0.310 | 0.163 | −0.108 | 0.180 | 0.069 | 0.059 |
TD | 0.359 | −0.322 | 0.254 | 0.258 | 0.063 | 0.198 |
TYE | 0.258 | 0.241 | -0.279 | 0.145 | 0.012 | 0.215 |
UAL | 0.125 | 0.373 | 0.530 | 0.177 | 0.000 | 0.019 |
FAL | 0.213 | −0.084 | -0.008 | 0.204 | 0.035 | 0.100 |
FeL | 0.185 | 0.027 | 0.143 | 0.211 | 0.028 | 0.006 |
TbL | 0.131 | 0.053 | −0.008 | 0.158 | 0.034 | 0.055 |
TbW | 0.161 | −0.094 | −0.082 | 0.280 | 0.023 | −0.091 |
FoL | 0.164 | 0.026 | 0.061 | 0.169 | 0.022 | 0.076 |
IMT | 0.271 | −0.457 | −0.078 | 0.413 | −0.466 | −0.724 |
Mount Ngoc Linh, on the northwestern border of the Kon Tum Massif, is the highest peak in central Vietnam at 2,598 m (
The new species has a restricted distribution in central Vietnam and Xekong Province, Lao PDR. A major threat to the new species in the area is habitat loss by agricultural extension for medicinal trees (e.g. Panax vietnamensis), illegal timber logging, and tourism development. In addition, the species Q. taoi is collected by local people for food. We suggest assessment of this species as Near Threatened in the IUCN Red List of Threatened Species because the continued survival of this species is largely dependent on the protection and rigorous management provided by local authorities of the protected areas in both countries.
In this study, we first uploaded to GenBank the 16S gene sequence of Quasipaa acanthophora from the type locality (Mau Son Mountain) in Lang Son Province, northern Vietnam. We confirm that Q. acanthophora is currently known only from Vietnam and does not correpond to a population of Q. spinosa according to
We are grateful to the directorates of the Ngoc Linh Nature Reserve, Forest Protection Department of Kon Tum Province for support of our field work. For the fruitful cooperation within joint research projects, we cordially thank S.V. Nguyen (IEBR, Hanoi), as well as T. Pagel and C. Landsberg (Cologne Zoo). We thank T.D. Tran (IEBR, Hanoi) for assistance in the field. We thank T.A. Tran (IEBR, Hanoi) for providing the map. We thank Annemarie Ohler and an anonymous reviewer for their helpful comments. This research is funded by the National Foundation for Science and Technology Development (NAFOSTED) under grant number 106.05-2020.02.
Specimens examined
Quasipaa acanthophora (n = 11): Vietnam: Lang Son Province: Cao Loc District: Mau Son Commune: IEBR A.5030–A.5035; Vietnam: Bac Giang Province: Son Dong District: Tay Yen Tu Nature Reserve: IEBR A.2013.62–A.2013.64, IEBR 3625–3626.
Quasipaa boulengeri (n = 13): Vietnam: Cao Bang Province: Nguyen Binh District: Phia Oac-Phia Den National Park: IEBR A.5007–A.5014, IEBR A.5039–A.5041; Vietnam: Ha Giang Province: Quan Ba District: IEBR A.5015–A.5016.
Quasipaa delacouri (n = 4): Vietnam: Tuyen Quang Province: Cham Chu Nature Reserve: IEBR A.5017; Vietnam: Ha Giang Province: Bac Me Nature Reserve: IEBR A.5018; Vietnam: Phu Tho Province: IEBR A.5019–A.5020.
Quasipaa verrucospinosa (n = 9): Vietnam: Vinh Phuc Province: Tam Dao National Park: IEBR A.5021; Vietnam: Ha Giang Province: Bac Me Nature Reserve: IEBR A.5022–A.5024; Vietnam: Tuyen Quang Province: Cham Chu Nature Reserve: IEBR A.5025–A.5027; Vietnam: Tuyen Quang Province: Na Hang District: IEBR A.5028–A.5029.