Gracixalus truongi
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Ordo: Anura
Familia: Rhacophoridae
Genus: Gracixalus
Name
Gracixalus truongi Tran & Pham & Le & Nguyen & Ziegler & Pham, 2023 sp. nov. – Wikispecies link – ZooBank link – Pensoft Profile
Material
Holotype: IEBR A.5004 (Field number TN 2020.09), adult male, collected by N.H. Nguyen, H.N. Tran, H.Q. Nguyen on 11 November 2020 in the karst forest in Ta Ma Commune (21°40'36.0"N, 103°31'96.7"E, at an elevation of 1,164 m asl.), Tuan Giao District, Dien Bien Province, Vietnam. Paratypes: IEBR A.5005 (Field number TN 2020.08), adult female, collected on 11 November 2020 (the same data as the holotype); IEBR A.5006 (Field number ĐB 2021.7), adult male, collected by H. Q. Nguyen & T. Q. Phan, on 30 December 2021, in Tuan Giao District, Dien Bien Province, Vietnam; ZVNU 09 (Field numbers Co9.16.24) and ZVNU 10 (Field Co9.16.36), two females, collected by A. V. Pham, N. B. Sung, L. M. Ha, T. Q. L. Hoang, and Q. T. Bui on 3 September 2016, in Long He Village (21°24'14.5"N, 103°28'41.5"E, at an elevation of 1,110 m asl.), Long He Commune, Thuan Chau District, Son La Province, Vietnam.
Diagnosis
The new species is assigned to the genus Gracixalus based on molecular phylogenetic analyses and the following morphological characters: the presence of intercalary cartilage between terminal and penultimate phalanges of digits, tips of digits expanded into large discs bearing circum-marginal grooves, the vomerine teeth being absent, horizontal pupil, tibia ~ 4–5× longer than wide, translucent skin, inner (first and second) and outer (third and fourth) fingers not opposable, and dorsum with an inversed Y-shaped dark brown marking (Fei et al. 2009[1]; Rowley et al. 2011[2], 2020[3]; Chen et al. 2018[4]; Yu et al. 2019[5]; Le et al. 2021[6]).
Gracixalus truongi sp. nov. is distinguishable from its congeners by a combination of the following morphological characters: (1) size relatively small, SVL 32.2–33.1 mm in males, 37.6–39.3 mm in females; (2) head slightly wider than long; (3) vomerine teeth absent; (4) snout round and long RL/SVL 0.17–0.19 in males, 0.16–0.17 in females; (5) spines on upper eyelid absent; (6) supratympanic fold distinct; (7) tympanum distinct; (8) dorsal skin smooth; (9) throat skin smooth and venter skin granular; (10) tibiotarsal projection absent; (11) webbing of fingers rudimentary, toes with moderately developed webbing; (12) dorsum moss-green, with an inverse Y-shaped dark green marking extended from interorbital region to posterior region of dorsum; (13) external vocal sac absent in males; (14) males with a nuptial pad on finger I.
Description of holotype
(male). Size small (SVL 33.1 mm), body robust, dorsoventrally compressed. Head slightly wider than long (HL 10.6 mm, HW 11.8 mm); snout round anteriorly in dorsal view, projecting beyond margin of the lower jaw; nostril round, without a lateral flap of skin, closer to tip of snout than to eye (NS 3.0 mm, EN 3.2 mm); canthus rostralis distinct and round; loreal region oblique and concave; rostral length greater than eye diameter (RL 4.8 mm, ED 4.5 mm); canthus rostralis round, loreal region oblique, concave; interorbital region flat, interorbital distance wider than internarial distance and upper eyelid width (IOD 3.9 mm, IND 3.7 mm, UEW 2.8 mm); distance between anterior corner of eyes (DAE 6.1 mm) ~ 57% distance between posterior corner of eyes (DPE 10.7 mm); pupil oval, horizontal; tympanum distinct (TYD 2.2 mm), round, half of the eye diameter but greater than tympanum-eye distance (TYE 1.5 mm); vomerine teeth absent; choanae small, oval; tongue cordate, deeply notched posteriorly; external vocal sacs absent.
Forelimbs robust; forearm and hand relative long (UAL/SVL 0.16), hand longer than forearm (FAL/SVL 0.45); relative finger lengths: I<II<IV<III; fingers webbing rudimentary; dermal ridge on sides of fingers absent; tips of all fingers with well-developed discs with distinct circum-marginal grooves, discs relatively wide compared to width of finger (fd3/fw3 1.9/1.2 mm), disc of finger III smaller than tympanum diameter; subarticular tubercles markedly elevated and prominent, round, one each on fingers I and II, two on fingers III and IV; nuptial pads prominent, oval; outer palmar tubercle divided into two.
Hindlimbs long (TbL/SVL 0.47, FoL/SVL 0.63); heels overlapping when held at right angles to the body; tibia length ~ 4× greater than tibia width (TbL/TbW 4.31), longer than thigh (FeL 15.1 mm) but shorter than foot length (FoL 20.8 mm); relative length of toes: I<II<III<V<IV; tips of all toes with well-developed discs with distinct circum-marginal grooves, discs slightly smaller than those of fingers; webbing formula I1–11/2II3/4–2III1–21/4IV2–1V; subarticular tubercles distinct, blunt, round: one on toes I and II, two on toes III and V, and three on toe IV; inner metatarsal tubercle small (IMT 1.3 mm); dermal ridge along outer side of tibia and tarsal fold absent; outer metatarsal and supernumerary tubercles absent; pointed projection at tibiotarsal articulation absent; tibio-tarsal articulation reaching between eye and nostril.
Skin texture: dorsal surface of head and body smooth; posterior part of tympanum, flank and lateral sides of limbs with small, flattened granules; spinules on upper eyelid absent; supratympanic fold distinct, extending from eye to angle of jaw; dorsolateral folds absent; throat and chest smooth, belly and ventral surface of thigh granular; dermal appendage at vent absent.
Coloration in life: background of dorsal surface of head, body and limbs moss-green with grey marking; with an inverse Y-shaped dark green marking, starting at the interorbital region, bifurcating into two branches on the shoulder, extending posteriorly; lateral side of body, dorsal surface of arms and limbs moss-green with dark green transverse bars; throat and chest white with dark brown marbling; belly immaculate white.
Coloration in preservative: Snout and dorsum grey with a dark brown pattern forming an inverse Y marking, notably a triangular pattern between eyes bifurcating into two bands continuing posteriorly; a dark pattern running from above cloaca forward to the middle of the back; lateral side of head and flank grey with dark spots; tympanum light brown; forelimb, dorsal surface of thigh, tibia and foot grey with some darker bands, posterior part of thigh below the vent yellowish brown with small white spots; throat and chest with dark brown marbling; belly immaculate cream to white; ventral part of forelimbs white; ventral surface of thighs white to grey; webbing grey.
Variation
Measurements and morphological characters of the type series are provided in Table 2 and photographs of the paratypes in life are presented in Fig. 4. Males are smaller than females (SVL 32.2–33.1 mm in males vs. 37.6–39.3 mm in females). The male specimens have a nuptial pad on finger I. The females contained yellowish cream eggs.
| Males | Females | ||||||
|---|---|---|---|---|---|---|---|
| Holotype | Paratype | Paratype | Paratype | Paratype | |||
| IEBR A.5004 | IEBR A.5006 | Min – Max | IEBR A.5005 | ZVNU 09 | ZVNU 10 | Min – Max | |
| SVL | 33.1 | 32.2 | 32.2–33.1 | 37.8 | 37.6 | 39.3 | 37.6–39.3 |
| HW | 11.8 | 13.0 | 11.8–13.0 | 14.6 | 14.6 | 14.8 | 14.6–14.8 |
| HL | 10.6 | 12.8 | 10.6–12.8 | 13.9 | 14.2 | 14.5 | 13.9–14.5 |
| MN | 9.5 | 10.8 | 9.5–10.8 | 11.9 | 12.1 | 12.9 | 11.9–12.9 |
| MFE | 7.4 | 8.2 | 7.4–8.2 | 9.3 | 9.2 | 9.9 | 9.2–9.9 |
| MBE | 4.1 | 5.0 | 4.1–5.0 | 4.9 | 5.1 | 5.5 | 4.9–5.5 |
| RL | 6.2 | 5.6 | 5.6–6.2 | 6.5 | 6.0 | 6.2 | 6.0–6.5 |
| ED | 4.5 | 5.1 | 4.5–5.1 | 5.1 | 5.1 | 5.3 | 5.1–5.3 |
| UEW | 2.8 | 3.2 | 2.8–3.2 | 3.7 | 3.5 | 3.9 | 3.5–3.9 |
| IND | 3.7 | 3.8 | 3.7–3.8 | 4.5 | 4.0 | 4.5 | 4.0–4.5 |
| IOD | 3.2 | 4.0 | 3.2–4.0 | 4.1 | 5.1 | 5.2 | 4.1–5.2 |
| DAE | 6.1 | 6.2 | 6.1–6.2 | 6.7 | 6.8 | 7.4 | 6.7–7.4 |
| DPE | 10.7 | 10.8 | 10.7–10.8 | 12.1 | 11.8 | 12.4 | 11.8–12.4 |
| NS | 3.0 | 2.6 | 2.6–3.0 | 3.0 | 2.8 | 2.9 | 2.8–3.0 |
| EN | 3.2 | 3.0 | 3.0–3.2 | 3.5 | 3.2 | 3.3 | 3.2–3.5 |
| TYD | 2.2 | 2.3 | 2.2–2.3 | 2.7 | 2.5 | 2.7 | 2.5–2.7 |
| TYE | 1.5 | 1.5 | 1.5–1.5 | 1.7 | 1.7 | 1.9 | 1.7–1.9 |
| UAL | 5.4 | 6.1 | 5.4–6.1 | 6.3 | 6.2 | 6.3 | 6.2–6.3 |
| FAL | 14.9 | 16.0 | 14.9–16.0 | 17.8 | 18.0 | 20.3 | 17.8–20.3 |
| NPL | 2.2 | 1.9 | 1.9–2.2 | ||||
| TFL | 8.3 | 8.9 | 8.3–8.9 | 8.9 | 8.9 | 10.2 | 8.9–10.2 |
| fd3 | 1.9 | 1.7 | 1.7–1.9 | 2.0 | 1.8 | 1.9 | 1.8–2.0 |
| fw3 | 1.2 | 1.0 | 1.0–1.2 | 1.3 | 1.2 | 1.3 | 1.2–1.3 |
| FeL | 15.1 | 15.2 | 15.1–15.2 | 17.7 | 17.1 | 20.2 | 17.1–20.2 |
| TbL | 15.5 | 18.0 | 15.5–18.0 | 18.1 | 19.2 | 21.9 | 18.1–21.9 |
| TbW | 3.6 | 4.0 | 3.6–4.0 | 4.2 | 4.0 | 4.8 | 4.0–4.8 |
| FoL | 20.8 | 23.2 | 20.8–23.2 | 24.2 | 25.1 | 28.3 | 24.2–28.3 |
| FTL | 12.6 | 13.0 | 12.6–13.0 | 14.8 | 15.4 | 16.7 | 14.8–16.7 |
| td4 | 1.8 | 1.6 | 1.6–1.8 | 1.9 | 1.7 | 1.8 | 1.7–1.9 |
| tw4 | 1.2 | 1.2 | 1.2–1.2 | 1.2 | 1.2 | 1.2 | 1.2–1.2 |
| IMT | 1.6 | 2.0 | 1.6–2.0 | 2.0 | 1.9 | 2.1 | 1.9–2.1 |
| RL/SVL | 0.19 | 0.17 | 0.17–0.19 | 0.17 | 0.16 | 0.16 | 0.16–0.17 |
| ED/RL | 0.73 | 0.91 | 0.73–0.91 | 0.78 | 0.85 | 0.85 | 0.78–0.85 |
| TYE/TYD | 0.68 | 0.65 | 0.65–0.68 | 0.63 | 0.68 | 0.70 | 0.63–0.70 |
| UAL/SVL | 0.16 | 0.19 | 0.16–0.19 | 0.17 | 0.16 | 0.16 | 0.16–0.17 |
| FAL/SVL | 0.45 | 0.50 | 0.45–0.50 | 0.47 | 0.48 | 0.52 | 0.47–0.52 |
| TbL/TbW | 4.31 | 4.50 | 4.31–4.50 | 4.31 | 4.80 | 4.56 | 4.31–4.80 |
| TbL/SVL | 0.47 | 0.56 | 0.47–0.56 | 0.48 | 0.51 | 0.56 | 0.48–0.56 |
| FoL/SVL | 0.63 | 0.72 | 0.63–0.72 | 0.64 | 0.67 | 0.72 | 0.64–0.72 |
| fd3/TYD | 0.86 | 0.74 | 0.74–0.86 | 0.74 | 0.72 | 0.70 | 0.70–0.74 |
Etymology
We name this new species in honor of our colleague, Prof. Dr. Truong Quang Nguyen from the Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, in recognition of his great contributions to the herpetofaunal exploration of the Indochina region. We recommend “Truong’s Treefrog” as the common English name of the new species and the common name in Vietnamese “Nhái cây trường”.
Ecological notes
The specimens were collected between 19:00 and 23:00 on a limestone cliff and on leaves, ~ 0.5–1.2 m above the ground. The surrounding habitat was secondary karst forest of medium and small hardwoods mixed with shrubs and vines. Air temperature was 13–18 °C and relative humidity was 65–80%. Other amphibian species found at the site were Leptobrachella sp., Kurixalus bisacculus (Taylor, 1962), Polypedates megacephalus Hallowell, 1861, and Rhacophorus orlovi (Ziegler & Köhler, 2001).
Distribution
Gracixalus truongi sp. nov. is currently known only from Dien Bien and Son La provinces, northwestern Vietnam (Fig. 5).
Comparisons
We compared the new species with other members of the genus Gracixalus and data obtained from the literature (Boulenger 1893[7]; Bourret 1937[8]; Hu et al. 1978[9]; Ye and Hu 1984[10]; Matsui and Orlov 2004[11]; Nguyen et al. 2008[12]; Rowley et al. 2011[2], 2014[13], 2020[3]; Mo et al. 2013[14]; Nguyen et al. 2013[15]; Matsui et al. 2015[16]; Matsui et al. 2017[17]; Zeng et al. 2017[18]; Chen et al. 2018[4]; Wang et al. 2018[19]; Yu et al. 2019[5]; Le et al. 2021[6]) (Table 3).
| Species | Adult male SVL (mm) | Adult female SVL (mm) | Conical tubercles on dorsum | Dorsal color in life | Vocal sac | Skin of body sides | Skin of throat | Finger webbing | Linea masculina | Tibiotarsal articulation |
| Gracixalus truongi sp. nov. | 32.2–33.1 | 37.6–39.3 | absent | moss green with grey | internal | smooth | smooth | absent | absent | reaching between eye and nostril |
| G. ananjevae | 20.0–32.0 | 43.4 | absent | ? | ? | coarsely granular | plain | rudimentary | ? | reaching eye |
| G. carinensis | 30.2–38.1 | ? | absent | purplish, reddish, or greyish brown | internal | ? | granular | rudimentary | ? | reaching eye |
| G. gracilipes | 20.0–24.0 | 26.4–28.8 | absent | greenish | internal | smooth with white stripe | smooth | rudimentary | ? | reaching eye |
| G. guangdongensis | 26.1–34.7 | 34.9–35.4 | absent | brown | ? | rough, black blotches | granular | absent | present | reaching between eye and nostril |
| G. jinggangensis | 27.9–33.8 | 31.6 | absent | brown to beige | ? | rough with tubercles | granular | rudimentary | ? | reaching eye |
| G. jinxiuensis | 23.5–26.3 | 29–30 | ? | brown | internal | rough with tubercles | granular | rudimentary | absent | reaching eye |
| G. lumarius | 38.9–41.6 | 36.3 | present | yellow | external | ? | granular | rudimentary | ? | ? |
| G. medogensis | 26.5 | ? | absent | grass green | internal | ? | granular | absent | present | reaching eye |
| G. nonggangensis | 27.1–35.3 | 26.8–27.3 | absent | yellowish-olive with dark-green mark | internal | rough with tubercles | granular | absent | absent | reaching tip of snout |
| G. quangi | 21.0–24.0 | 26.8–27.3 | present, small | olive-green | external | with black blotches | smooth | absent | ? | ? |
| G. quyeti | 28.5 | 34.0 | present | brownish to moss-green | ? | rough with sharp tubercles | smooth | rudimentary | ? | reaching to snout |
| G. sapaensis | 20.8–29.6 | 27.2–39.5 | absent | Golden ochre | ? | coarsely scattered with large tubercles | ? | rudimentary | ? | reaching eye |
| G. seesom | 21.6–23.0 | 23.2–25.4 | absent | tan | external | with large tubercles and white blotches | smooth | rudimentary | ? | reaching between eye and nostril |
| G. supercornutus | 22.0–24.1 | ? | present, bigger horn-like | green with brown spots | ? | ? | granular | ? | ? | ? |
| G. tianlinensis | 30.3–35.9 | 35.6–38.7 | absent | brown to beige | external | ? | granular | absent | ? | ? |
| G. trieng | 37.2–41.4 | present | brown or yellowish | present | ? | granular | rudimentary | ? | ? | |
| G. yunnanensis | 26.0–34.2 | ? | present, small | yellow brown or red brown | external | smooth, no black blotches | granular | rudimentary | present | reaching eye |
| G. ziegleri | 28.1–30.0 | 36.7–41.2 | present | yellowish brown | internal | rough, black blotches | granular | rudimentary | absent | reaching tip of snout |
| Species | Snout | White patch on temporal region | Tibiotarsal projection | Supratympanic fold | Venter | Nuptial pads | Heels | Iris | Linea masculina | Tibiotarsal articulation |
| Gracixalus truongi sp. nov. | rounded | absent | absent | distinct | immaculate white | on finger I | overlapping | brown and moss green | absent | reaching between eye and nostril |
| G. ananjevae | slightly pointed | absent | absent | distinct | immaculate | on finger I | overlapping | ? | ? | reaching eye |
| G. carinensis | round | absent | absent | distinct | immaculate white | ? | ? | ? | ? | reaching eye |
| G. gracilipes | triangularly pointed | absent | absent | distinct | yellowish white | on fingers I and II | overlapping | brown | ? | reaching eye |
| G. guangdongensis | triangularly pointed | present | present | distinct | throat and chest creamy white, belly light brown, semi-transparent | on finger I | overlapping | brown | present | reaching between eye and nostril |
| G. jinggangensis | triangularly pointed | absent | absent | distinct | Throat and chest dirty white with dark specks, belly white anteriorly with dark marking and posteriorly yellowish, semi-transparent | on fingers I and II | just meeting | golden | ? | reaching eye |
| G. jinxiuensis | round | absent | absent | distinct | gray-brown with dark marbling | on finger I | just meeting | pale brown | absent | reaching eye |
| G. lumarius | round | absent | absent | indistinct | opaque pink | on finger I | ? | dark gold | ? | ? |
| G. medogensis | round | absent | absent | distinct | pale green | on finger I | overlapping | ? | present | reaching eye |
| G. nonggangensis | round | absent | absent | distinct | white with dark marbling, semi- transparent | on finger I | overlapping | olive | absent | reaching tip of snout |
| G. quangi | triangularly pointed | present | present | distinct | opaque white with translucent pale green margins | on finger I | ? | bronze | ? | ? |
| G. quyeti | round | absent | absent | indistinct | belly immaculate white | ? | overlapping | yellow moss green | ? | reaching to snout |
| G. sapaensis | round | absent | absent | distinct | throat, chest, and belly light yellow, with dark marking | on finger I | overlapping | golden | ? | reaching eye |
| G. seesom | triangularly pointed | absent | absent | distinct | anterior belly opaque white and posterior belly translucent | absent | overlapping | golden | ? | reaching between eye and nostril |
| G. supercornutus | pointed | present | present | distinct | light with white spots | ? | ? | Pale yellow | ? | ? |
| G. tianlinensis | round | absent | absent | distinct | throat and chest gray with dark specks, belly creamy white, opaque | on fingers I and II | ? | bronze | ? | ? |
| G. trieng | rounded | ? | absent | distinct | throat and chest mostly yellowish brown, with dark mottling; belly pinkish brown | on fingers I and II | ? | pale gold | ? | ? |
| G. yunnanensis | round | absent | absent | distinct | orangish with yellow spots, immaculate, semi-transparent | on finger I | overlapping | bronze | present | reaching eye |
| G. ziegleri | triangularly pointed | absent | absent | distinct | throat and chest dirty white with moderate dark specks, belly white cream with large dark blotches, | on finger I | overlapping | golden | absent | reaching tip of snout |
In terms of dorsal color pattern Gracixalus truongi sp. nov. is similar to Theloderma annae Nguyen, Pham, Nguyen, Ngo & Ziegler, 2016 from Hoa Binh anh Ninh Binh provinces. In addition, Gracixalus truongi sp. nov. and Theloderma annae also have similar life histories, both inhabiting limestone karst forest far from water sources. However, Gracixalus truongi differs from Theloderma annae by a larger size (SVL 32.1–33.1 in males, 37.6–39.3 mm in females vs. 27.1–28.5 mm in males, 30.3–32.6 mm in females), the presence of a dark inverse Y-marking on dorsum (vs. absent), and a higher ratio of TYD/TYD (0.67 in males and 0.67 in females vs. 0.39 in males and 0.32 in females) (Nguyen et al. 2016[20]).
Original Description
- Tran, T; Pham, A; Le, M; Nguyen, N; Ziegler, T; Pham, C; 2023: A new species of Gracixalus (Anura, Rhacophoridae) from northwestern Vietnam ZooKeys, 1153: 15-35. doi
Images
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Other References
- ↑ Fei L, Hu S, Ye C, Huang Y (2009) Fauna Sinica. Amphibia (Vol. 2).Anura, Science Press, Beijing, 887 pp.
- ↑ 2.0 2.1 2.2 Rowley J, Dau Q, Nguyen T, Cao T, Nguyen S (2011) A new species of Gracixalus (Anura: Rhacophoridae) with a hyperextended vocal repertoire from Vietnam.Zootaxa3125: 22–38. https://doi.org/10.1163/15685381-00003007
- ↑ 3.0 3.1 3.2 Rowley J, Le D, Hoang H, Cao T, Dau Q (2020) A new species of phytotelm breeding frog (Anura: Rhacophoridae) from the Central Highlands of Vietnam.Zootaxa4779: 341–354. https://doi.org/10.11646/zootaxa.4779.3.3
- ↑ 4.0 4.1 4.2 Chen W, Bei Y, Liao X, Zhou S, Mo Y (2018) A new species of Gracixalus (Anura: Rhacophoridae) from West Guangxi, China.Asian Herpetological Research9: 74–84. https://doi.Org/10.11646/zootaxa.3616.1.5
- ↑ 5.0 5.1 5.2 Yu G, Hui H, Wang J, Rao D, Wu Z, Yang J (2019) A new species of Gracixalus (Anura, Rhacophoridae) from Yunnan, China.ZooKeys851: 91–111. https://doi.org/10.3897/zookeys.851.32157
- ↑ 6.0 6.1 6.2 Le D, Do Y, Tran T, Nguyen T, Orlov N, Ninh T, Nguyen T (2021) A new species of Gracixalus (Anura: Rhacophoridae) from northern Vietnam.Russian Journal of Herpetology28(3): 111–122. https://doi.org/10.30906/1026-2296-2021-28-3-111-122
- ↑ 7.0 7.1 Boulenger G (1893) Concluding report on the reptiles and batrachians obtained in Burma by Signor L. Fea dealing with the collection made in Pegu and the Karin Hills in 1887–88.Annali del Museo Civico di Storia Naturale di Genova13: 304–347. https://doi.Org/10.5962/bhl.part.9543
- ↑ 8.0 8.1 Bourret R (1937) Notes herpétologiques sur l’Indochine française. XIV. Les batraciens de la collection du Laboratoire des Sciences Naturelles de l’Université. Descriptions de quinze espèces ou variétés nouvelles.Annexe au Bulletin Général de l’Instruction Publique1937: 5–56. https://doi.org/10.5962/bhl.part.22065
- ↑ 9.0 9.1 Hu S, Fei L, Ye C (1978) Three new amphibian species in China. Materials for Herpetological Research 4: e20.
- ↑ 10.0 10.1 Ye C, Hu S (1984) A new species of Philautus (Anura: Rhacophoridae) from Xizang Autonomous Region.Acta Herpetologica Sinica3: 67–69.
- ↑ 11.0 11.1 Matsui M, Orlov N (2004) A new species of Chirixalus from Vietnam (Anura: Rhacophoridae).Zoological Science21(6): 671–676. https://doi.org/10.2108/zsj.21.671
- ↑ 12.0 12.1 Nguyen T, Hendrix R, Böhme W, Vu T, Ziegler T (2008) A new species of the genus Philautus (Amphibia: Anura: Rhacophoridae) from the Truong Son Range, Quang Binh Province, central Vietnam.Zootaxa1925(1): 1–13. https://doi.org/10.11646/zootaxa.1925.1.1
- ↑ 13.0 13.1 Rowley J, Le D, Dau V, Hoang H, Cao T (2014) A striking new species of phytotelm breeding tree frog (Anura: Rhacophoridae) from central Vietnam.Zootaxa3785(1): 25–37. https://doi.org/10.11646/zootaxa.3785.1.2
- ↑ 14.0 14.1 Mo Y, Zhang W, Luo Y, Zhou S, Chen W (2013) A new species of the genus Gracixalus (Amphibia: Anura: Rhacophoridae) from Southern Guangxi, China.Zootaxa3616(1): 61–72. https://doi.org/10.11646/zootaxa.3616.1.5
- ↑ 15.0 15.1 Nguyen T, Le M, Pham C, Nguyen T, Bonkowski M, Ziegler T (2013) A new species of Gracixalus (Amphibia: Anura: Rhacophoridae) from northern Vietnam.Organisms, Diversity & Evolution13(2): 203–214. https://doi.org/10.1007/s13127-012-0116-0
- ↑ 16.0 16.1 Matsui M, Khonosue W, Panha S, Eto K (2015) A new tree frog of the genus Gracixalus from Thailand (Amphibia: Rhacophoridae).Zoological Science32(2): 204–210. https://doi.org/10.2108/zs140238
- ↑ 17.0 17.1 Matsui M, Ohler A, Eto K, Nguyen T (2017) Distinction of Gracixalus carinensis from Vietnam and Myanmar, with description of a new species.Alytes33: 25–37.
- ↑ 18.0 18.1 Zeng Z, Zhao J, Chen C, Chen G, Zhang Z, Wang Y (2017) A new species of the genus Gracixalus (Amphibia: Anura: Rhacophoridae) from Mount Jinggang, southeastern China.Zootaxa4250(2): 171–185. https://doi.org/10.11646/zootaxa.4250.2.3
- ↑ 19.0 19.1 Wang J, Zeng Z, Lyu Z, Liu Z, Wang Y (2018) Description of a new species of Gracixalus (Amphibia: Anura: Rhacophoridae) from Guangdong Province, southeastern China.Zootaxa4420(2): 251–269. https://doi.org/10.11646/zootaxa.4420.2.7
- ↑ Nguyen T, Pham T, Nguyen T, Ngo T, Ziegler T (2016) A new species of Theloderma (Amphibia: Anura: Rhacophoridae) from Vietnam.Zootaxa4168(1): 171–186. https://doi.org/10.11646/zootaxa.4168.1.10
