Research Article |
Corresponding author: Guohua Yu ( yugh2018@126.com ) Corresponding author: Zhengjun Wu ( wu_zhengjun@aliyun.com ) Academic editor: Annemarie Ohler
© 2023 Junkai Huang, Xiao Long Liu, Lingyun Du, Justin M. Bernstein, Shuo Liu, Yun Yang, Guohua Yu, Zhengjun Wu.
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:
Huang J, Liu XL, Du L, Bernstein JM, Liu S, Yang Y, Yu G, Wu Z (2023) A new species of Bush frog (Anura, Rhacophoridae, Raorchestes) from southeastern Yunnan, China. ZooKeys 1151: 47-65. https://doi.org/10.3897/zookeys.1151.95616
|
In this study, based on morphological and molecular data, a new bush frog species is described from Yunnan, China. Eleven samples of Raorchestes malipoensis sp. nov. were collected from Malipo County, southeastern Yunnan. This species can be distinguished from other congeners by a combination of 13 morphological characters. Phylogenetic analyses based on the 16S rRNA gene indicate that these individuals form a monophyletic group, and genetic divergence between this clade and its closest relatives is higher than 3.1%, which is comparable to the divergence between recognized Raorchestes species. The discovery of this new species suggests that additional extensive surveys in the southeastern Yunnan would yield more amphibian lineages yet unknown to science.
Kurixalus gryllus, Raorchestes gryllus, Raorchestes malipoensis sp. nov., taxonomy
The genus Raorchestes Biju, Shouche, Dubois, Dutta & Bossuyt, 2010 belongs to the family Rhacophoridae Hoffman, 1932. It includes bush frogs with adult size ranging from 10.0 mm to 50.5 mm (
Many Raorchestes species from the region were described with few diagnostic characters and limited morphological data, which hampers the identification of these small-sized bush frogs (
In this work we studied specimens allocated to Raorchestes from Malipo County. This county is located in the southeast of Yunnan Province, and lies on the China-Vietnam border where few herpetological investigations have been conducted. During the fieldwork, we collected 11 specimens of a small-sized bush frog that could be assigned to the genus Raorchestes based on morphological and molecular evidence. Phylogenetically, these specimens were grouped together with a misidentified “R. gryllus” from Pac Ban, Tuyen Quang, northern Vietnam. However, considering that the type locality of Philautus gryllus, Langbian Plateau, is 1200 km far from the China-Vietnam border and that obvious morphological differences exist between Philautus gryllus and the lineage consisting of individuals from China-Vietnam border region, we consider that these specimens represent a new species that we formally describe here.
Fieldwork was conducted at Malipo County, Yunnan Province, China (23.182°N, 104.78°E, elevation 1496 m). Six specimens were collected on 7 May 2019 (Figs
All the measurements were made with slide calipers to the nearest 0.1 mm. Morphological terminology and measurement methods followed
Measurements (mm) of adult specimens in the type series of Raorchestes malipoensis sp. nov. Abbreviations defined in the Materials and methods.
Sex | Males (n = 4) | Females (n = 2) | Males (n = 3) | Females (n = 2) | |||||||
---|---|---|---|---|---|---|---|---|---|---|---|
Catalog No. | SWFU 3110 | SWFU 3113 | SWFU 3114 | SWFU 3116 | SWFU 3111 | SWFU 3112 |
|
|
|
|
|
SVL | 17.1 | 17.0 | 16.5 | 14.7 | 19.3 | 19.0 | 17.5 | 17.7 | 17.3 | 18.7 | 18.3 |
HL | 5.7 | 5.8 | 6.3 | 5.2 | 6.5 | 7.9 | 5.9 | 6.4 | 5.7 | 6.3 | 6.7 |
HW | 7.7 | 7.5 | 8.2 | 5.5 | 8.2 | 7.9 | 6.5 | 6.8 | 6.4 | 7 | 7 |
SL | 2.5 | 2.6 | 1.8 | 2.5 | 2.9 | 2.6 | 2 | 2.6 | 2.4 | 2.5 | 2.2 |
INS | 1.5 | 2.2 | 2.2 | 2.0 | 2.2 | 2.1 | 2.2 | 2.1 | 2.3 | 2.2 | 2 |
IOS | 2.7 | 2.7 | 2.9 | 2.9 | 3.2 | 2.9 | 2.9 | 2.6 | 2.9 | 2.7 | 2.9 |
UEW | 1.2 | 1.2 | 1.9 | 1.4 | 1.6 | 1.7 | 1.6 | 1.3 | 1.4 | 1.4 | 1.7 |
EHD | 2.2 | 2.1 | 2.1 | 2.6 | 2.6 | 2.8 | 2.4 | 2.6 | 2.4 | 2.5 | 2.2 |
TD | 1.4 | 1.4 | 1.5 | 1.3 | 1.1 | 1.5 | 1.2 | 1.5 | 1.4 | 1.6 | 1.4 |
FAHL | 9.3 | 9.3 | 8.8 | 7.0 | 8.6 | 9.6 | 7.8 | 8.2 | 7.2 | 7.3 | 7.1 |
HAL | 5.6 | 5.6 | 5.3 | 4.2 | 5.2 | 5.6 | 5.3 | 5.2 | 5.3 | 4.6 | 5.4 |
LAW | 1.7 | 2.0 | 1.6 | 1.3 | 1.4 | 1.6 | 1.5 | 1.2 | 1.6 | 1.3 | 1.3 |
TBL | 9.1 | 8.7 | 8.4 | 7.5 | 9.1 | 9.2 | 9 | 8.8 | 8 | 8.7 | 8.8 |
FML | 7.5 | 9.2 | 8.3 | 7.1 | 8.0 | 10.2 | 7.5 | 8 | 7.5 | 8.2 | 7.9 |
TBW | 2.1 | 2.6 | 1.8 | 1.5 | 2.0 | 2.8 | 1.7 | 1.7 | 1.8 | 1.8 | 1.9 |
TFL | 9.8 | 10.9 | 10.6 | 8.8 | 10.0 | 11.8 | 10.8 | 11 | 10.1 | 9.6 | 10.7 |
The source of morphological data for Raorchestes species used in this study.
ID | Raorchestes species | Literature |
---|---|---|
3 | Raorchestes ghatei Padhye, Sayyed, Jadhav, & Dahanukar, 2013 |
|
4 | Raorchestes parvulus (Boulenger, 1893) |
|
5 | Raorchestes cangyuanensis Wu, Suwannapoom, Xu, Murphy, & Che, 2019 |
|
6 | Raorchestes longchuanensis (Yang & Li, 1978) |
|
7 | Raorchestes menglaensis (Kou, 1990) |
|
8 | Raorchestes hillisi Jiang Ren, Guo, Wang & Li, 2020 |
|
9 | Raorchestes huanglianshan Jiang, Wang, Ren, & Li, 2020 |
|
10 | Raorchestes dulongensis Wu, Liu, Gao, Wang, Li, Zhou, Yuan, & Che, 2021 |
|
11 | Raorchestes andersoni (Ahl, 1927) |
|
12 | Raorchestes rezakhani Al-Razi, Maria, & Muzaffar, 2020 |
|
13 | Raorchestes annandalii (Boulenger, 1906) |
|
Total DNA was extracted using a commercial tissue DNA isolation kit (Chenlu Biotech, China). For seven specimens in this study, the mitochondrial gene 16S ribosomal RNA (16S rRNA) gene was sequenced. The fragments of 16S rRNA were amplified using primers 16Sar-L (5’–CGCCTGTTTATCAAAAACAT–3’) and 16Sbr-H (5’–CCGGTCTGAACTCAGATCACGT–3’) (
Information on voucher numbers, GenBank accession numbers, and localities of specimens used in this study; for collections and their abbreviations see Material and methods.
Species | Voucher No. | GenBank No. | Locality | Resource |
---|---|---|---|---|
Ingroup | ||||
Raorchestes malipoensis sp. nov. | SWFU 3110 | ON128247 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. | SWFU 3111 | ON128241 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. | ROM 30288 | GQ285674 | Pac Ban, Tuyen Quang, Vietnam |
|
Raorchestes malipoensis sp. nov. |
|
ON128246 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. |
|
ON128245 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. |
|
ON128244 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. |
|
ON128243 | Malipo, Yunnan, China | This study |
Raorchestes malipoensis sp. nov. |
|
ON128242 | Malipo, Yunnan, China | This study |
Raorchestes dulongensis | KIZ 035082 | MW537814 | Qinlangdang, Yunnan, China |
|
Raorchestes hillisi | CIB 116331 | MT488411 | Xiding, Yunnan, China |
|
Raorchestes longchuanensis | KIZ 048468 | MN475870 | Unknown |
|
Raorchestes parvulus | LSUHC:11118 | MH590201 | Gunung Stong, Kelantan, Malaysia |
|
Raorchestes menglaensis | CIB 116349 | MT488410 | Menglun, Yunnan, China |
|
Raorchestes huanglianshan | CIB 116365 | MT488414 | Lvchun, Yunnan, China |
|
Raorchestes cangyuanensis | KIZ 015855 | MN475866 | Cangyuanensis, Yunnan, China |
|
Raorchestes tuberohumerus | CESF 148 | JX092697 | Western Ghats, India |
|
Raorchestes bombayensis | CESF 1010 | JX092657 | Western Ghats, India |
|
Raorchestes ghatei | AGCZRL Amphibia 128 | KF366391 | Western Ghats, India |
|
Raorchestes griet | CESF 073 | JX092654 | Western Ghats, India |
|
Raorchestes coonoorensis | CESF 439 | JX092716 | Western Ghats, India |
|
Raorchestes charius | CESF 132 | JX092691 | Western Ghats, India |
|
Raorchestes marki | CESF 467 | JX092719 | Western Ghats, India |
|
Raorchestes indigo | CESF 138 | KM596557 | Kudremukh Massif, Western Ghats, India |
|
Raorchestes emeraldi | CESF 1365 | KM596556 | Valparai plateau, Western Ghats, India |
|
Raorchestes ponmudi | CESF 063 | JX092651 | Western Ghats, India |
|
Raorchestes aureus | CESF 1164 | KM596540 | Malabar, Western Ghats, India |
|
Raorchestes montanus | CESF 130 | KM596552 | Western Ghats, India |
|
Raorchestes tinniens | CESF 438 | JX092715 | Western Ghats, India |
|
Raorchestes primarrumfi | CESF 442 | KM596575 | Nilgiri Massif, Western Ghats,India |
|
Raorchestes signatus | Unknow | AY141841 | Sri Lanka |
|
Raorchestes chromasynchysi | CESF 1127 | JX092667 | Western Ghats, India |
|
Raorchestes chotta | CESF 1003 | JX092656 | Western Ghats, India |
|
Raorchestes nerostagona | CESF 1061 | JX092661 | Western Ghats, India |
|
Raorchestes kadalarensis | CESF 1766 | JX092701 | Western Ghats, India |
|
Raorchestes agasthyaensis | CESF 492 | JX092723 | Western Ghats, India |
|
Raorchestes travancoricus | CESF 473 | JX092721 | Western Ghats, India |
|
Raorchestes luteolus | CESF 1012 | JX092659 | Western Ghats, India |
|
Raorchestes beddomii | CESF 072 | JX092653 | Western Ghats, India |
|
Raorchestes theuerkaufi | CESF 1342 | JX092693 | Western Ghats, India |
|
Raorchestes munnarensis | CESF 094 | JX092655 | Western Ghats, India |
|
Raorchestes anili | CESF 386 | JX092708 | Western Ghats, India |
|
Raorchestes resplendens | CESF 1258 | JX092683 | Western Ghats, India |
|
Raorchestes dubois | CESF 114 | JX092668 | Western Ghats, India |
|
Raorchestes kakachi | CESF 1385 | KM596558 | Western Ghats, India |
|
Raorchestes kaikatti | CESF 444 | JX092718 | Western Ghats, India |
|
Raorchestes sushili | CESF 1259 | JX092684 | Western Ghats, India |
|
Raorchestes flaviocularis | CESF 1252 | KM596549 | Manalar Plateau, Western Ghats, India |
|
Raorchestes ochlandrae | CESF 1111 | JX092666 | Western Ghats, India |
|
Raorchestes manohari | CESF 1187 | JX092674 | Western Ghats, India |
|
Raorchestes uthamani | CESF 483 | JX092722 | Western Ghats, India |
|
Raorchestes chlorosomma | FB-2008c | EU450017 | Munnar, Idukki, Kerala, India |
|
Raorchestes crustai | CESF 1199 | JX092677 | Western Ghats, India |
|
Raorchestes graminirupes | CESF 044 | JX092649 | Western Ghats, India |
|
Raorchestes johnceei | CESF 1236 | JX092679 | Western Ghats, India |
|
Raorchestes glandulosus | CESF 1080 | JX092665 | Western Ghats, India |
|
Raorchestes jayarami | CESF 1260 | JX092686 | Western Ghats, India |
|
Raorchestes bobingeri | CESF 1238 | JX092680 | Western Ghats, India |
|
Raorchestes akroparallagi | CESF 061 | JX092650 | Western Ghats, India |
|
Outgroup | ||||
Philautus abditus | ROM 33145 | GQ285673 | Krong Pa, Gia Lai, Vietnam |
|
The final DNA sequence dataset is consisted of 59 sequences and the length of the sequence alignment is 542 base pairs (bp) (Table
Uncorrected pairwise sequence divergence (%) among 16S ribosomal RNA mtDNA sequences, including R. malipoensis sp. nov., R. dulongensis, R. hillisi, R. longchuanensis, R. menglaensis, R. huanglianshan, R. cangyuanensis, R. parvulus, R. bombayensis, R. tuberohumerus, and R. ghatei as shown in phylogenetic tree presented in Fig.
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 |
---|---|---|---|---|---|---|---|---|---|---|
R. malipoensis sp. nov. | ||||||||||
R. bombayensis | 4.5 | |||||||||
R. tuberohumerus | 4.8 | 2.0 | ||||||||
R. ghatei | 4.5 | 4.2 | 4.0 | |||||||
R. parvulus | 5.9 | 5.1 | 5.9 | 5.4 | ||||||
R. cangyuanensis | 5.7 | 6.6 | 6.8 | 5.6 | 7.1 | |||||
R. longchuanensis | 3.1 | 4.7 | 4.9 | 4.0 | 4.0 | 5.6 | ||||
R. menglaensis | 5.2 | 4.2 | 4.9 | 5.7 | 2.0 | 5.9 | 4.7 | |||
R. hillisi | 4.3 | 4.9 | 5.2 | 4.3 | 61 | 6.3 | 4.2 | 5.4 | ||
R. huanglianshan | 6.0 | 6.1 | 6.4 | 5.2 | 45 | 6.8 | 5.4 | 4.5 | 5.4 | |
R. dulongensis | 5.2 | 6.1 | 6.4 | 5.0 | 6.6 | 6.6 | 4.2 | 6.2 | 3.3 | 5.7 |
Phylogram of Raorchestes derived from analyses of concatenated DNA fragments of the mitochondrial 16S rRNA. Nodal support values with Bayesian posterior probability (BPP) > 0.95 / ML inferences (ML-BS) > 70 are shown near the respective nodes. A “-” denotes a Bayesian posterior probability < 0.95 and bootstrap support < 70. The scale bar represents 0.1 nucleotide substitutions per site.
“Pseudophilautus gryllus” (
“Raorchestes gryllus” (
SWFU 3110, SWFU 3113, SWFU 3114, SWFU 3116,
The genus Raorchestes is a group of small frogs, diagnosed primarily on the basis of an adult snout-vent length between 15 and 45 mm; vomerine teeth absent; large gular pouch transparent while calling; nocturnally active; direct development without free-swimming tadpoles in all species for which the development is known (
Adult male (Fig.
Forelimbs fairly robust (FAHL 8.2 mm); relative finger lengths: I < II < IV < III, tips of all four fingers expanded into discs with circum-marginal grooves; all fingers with lateral dermal fringes on both sides; subarticular tubercles distinct, rounded; supernumerary tubercles absent; no webbing between fingers; inner and outer metacarpal tubercle indistinct; nuptial pad is small and milky white on dorsal surface of the first finger.
Foot long and relatively robust (TFL 11 mm), longer than tibia length (TBL 8.8 mm); relative toe lengths: I < II < V < III < IV; tips of toes with discs having circum-marginal grooves, toe discs smaller than finger discs; all toes with lateral dermal fringes on both sides; subarticular tubercles distinct, rounded; supernumerary tubercles absent; webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); inner metatarsal tubercle rounded, outer metatarsal tubercle absent.
Dorsal surfaces of head, body, forelimbs, thighs, and tibia rough with small granules; upper eyelid with several small granules; throat, chest, and ventral surfaces of forelimbs smooth; abdomen, ventral side of thigh, and area around vent with granules; dorsolateral folds absent.
For coloration of the holotype in life see Fig.
After preservation in alcohol, the general pattern did not change. Dorsal color changed to grayish brown, the blotches or spots blackish brown, discs on the fingers become pale gray similar to the body color, ventral side become whiter (Fig.
The specific epithet is named for the type locality, Malipo County, Yunnan Province, China. We suggest “Malipo Bush Frog” as its English common name, and “Ma Li Po Guan Shu Wa (麻栗坡灌树蛙)” as its Chinese common name.
Currently known from the type locality, Malipo County (Fig.
The measurements are given in Table
Rather than comparing R. malipoensis sp. nov. to all known Raorchestes, we focus on our morphological comparison with phylogenetically closely related taxa and species without genetic data in adjacent countries (Table
Comparison of R. malipoensis sp. nov. with phylogenetically closely related taxa or those with no genetic data in surrounding countries. “–” means unknown.
Species | R. malipoensis sp. nov. | R. menglaensis | R. parvulus | R. dulongensis | R. hillisi | R. huanglianshan | R. cangyuanensis | R. ghatei | R. rezakhani | R. annandalii | R. bombayensis | R. tuberohumerus | R. longchuanensis | R. andersoni |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SVL of adult males (in mm) | 14.6–17.7, n = 7 | 16.6–21.6, n = 14 | – | 15.0–19.0, n = 3 | 14.5–17.7, n = 3 | 17.0–19.6, n = 11 | 16.1–19.0 mm, n = 3 | 19.1–25.5, n = 9 | 18.8–19.0 mm, n = 4 | – | 30 mm, n = – | 17.4–18.2 mm, n = 3 | 21.4–23.9 mm, n = 5 | 13.5–24.0 mm, n = 2 |
SVL of adult females (in mm) | 18.3–19.3, n = 4 | 18.9–20.5, n = 2 | 23.6, n = 1 | – | 17.5, n = 1 | 21.5, n = 1 | – | 15.4–29.8, n = 13 | – | 17.0 mm, n = 1 | – | – | – | |
SVL of adult (in mm) | 14.6–19.3, n = 11 | 16.6–21.6, n = 18 | 23.6, n = 1 | 15.0–19.0, n = 3 | 14.5–17.7, n = 4 | 17.0–21.5, n = 4 | 16.1–19.0 mm, n = 3 | 15.4–29.8, n = 22 | 18.8–19.0 mm, n = 4 | 17.0 mm, n = 1 | 30 mm, n = – | 17.4–18.2 mm, n = 3 | 21.4–23.9 mm, n = 5 | 13.5–24.0 mm, n = 2 |
IOS/EHD | IOS > EHD, or IOS = EHD | IOS > EHD | IOS < EHD | IOS < EHD | IOS < EHD | IOS > EHD, or IOS=EHD | IOS < EHD | – | IOS < EHD | IOS > EHD, or IOS = EHD | – | IOS > EHD, or IOS = EHD | IOS > EHD | IOS < EHD |
HDW/HDL | HDW > HDL | HDW < HDL | HDW > HDL | HDW < HDL | HDW < HDL | HDW > HDL | HDW > HDL | HDW > HDL | HDW > HDL | HDW < HDL | – | HDW > HDL | HDW ≈ HDL | HDW > HDL |
Tympanum | Distinct | Indistinct | Distinct | Distinct | Distinct | Distinct | Indistinct | Indistinct | Indistinct | Distinct | Indistinct | Indistinct | Distinct | Distinct |
Nuptial pad | Small and milky white | white nuptial pad | – | Absent | Present | Present | Reddish nuptial pad | Absent | Absent | – | – | – | Present | – |
Toe web | I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V | II 1 – 2 III 1 – 21/2 IV 21/2 – 1 V | Webbing present, medium | Rudimentary web | II 1 – 2 III 1–21/2 IV 21/2 – 1 V | II 1 – 2 III 1 – 2 – IV 2 – 1 V | Rudimentary web | I 2 – 2 II 2 – 2½ III 2 – 3 IV 2½ – 2 V | I2 – 2 II 1¾ – 2 III 1½ – 3 IV 2¾ – 2 V | Rudimentary web | 1/3 webbing | Rudimentary web | 1/4 webbing | 1/3 webbing |
Lateral dermal fringe | Present | Absent | – | – | Present | Absent | Present | Present | Absent | Present | – | – | – | – |
Disc color | Yellow | Not orange in life | – | Greyish or orange | – | Orange | Orange | – | Reddish or whitish | – | – | – | Reddish, orange, or whitish | orange |
Inner metacarpal tubercle | Indistinct | Present | Present | Present | Indistinct | Indistinct | – | – | Absent | Present | – | – | Present | Present |
Outer metacarpal tubercle | Indistinct | Present | Present | Present | Indistinct | Indistinct | – | – | Absent | Present | – | – | Present | Present |
Inner metatarsal tubercle | Round | Present | Present | Round | Round | Round | Round | Round | Absent | Absent | – | Present | Present | Present |
Outer metatarsal tubercle | Absent | Present | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | – | Absent | Absent | Absent |
Relative toe lengths | I < II < V < III < IV | III ≈ V, or V > III | I < II < V < III < IV | I < II < V < III < IV | I<II<III<V<IV | I<II<III<V<IV | I<II<V<III<IV | I<II<V=III<IV | I < II < V < III < IV | I < II < V = III < IV | – | I < II ≤ V < III < IV | III ≈ V | I < II < III = V < IV |
Range | Malipo, Yunnan, China and the north of Vietnam | Mengla, Yunnan, China | Indochina Peninsula and peninsular Malaysia | Gongshan, Yunnan, China | Menghai, Yunnan, China | Lvchun, Yunnan, China | Cangyuan, Yunnan, China | Western Ghats, India | Northeastern Bangladesh | Himalayas and northeastern India | Western Ghats, India | Western Ghats, India | Yunnan, China and Lai Chau,Vietnam | India, North Myanmar, Tibet and Yunnan, China |
The new species differs from R. menglaensis by 1) tubercles absent along the outer side of the forearm and foot; (2) head wider than long; (3) tympanum distinct (TD 1.1–1.6 mm, n = 11); (4) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); (5) lateral dermal fringe present (6) inner and outer metacarpal tubercle indistinct; (7) outer metatarsal tubercle absent; and (8) relative toe lengths: I < II < V < III < IV (vs. a series of tubercles along the outer side of the forearm and foot; head length and head width are approximately the same; tympanum indistinct; webbing formula (II 1 – 2 III 1 – 21/2 IV 21/2–1 V); lateral dermal fringe present; inner and outer metatarsal tubercle present; outer metatarsal tubercle present; relative toe lengths: III ≈ V, or V > III).
The new species differs from R. parvulus by (1) smaller female body size (females 18.3–19.3 mm, n = 4); (2) interorbital distance larger than eye horizontal diameter; and (3) inner and outer metacarpal tubercle indistinct; (vs. female 23.6 mm, n = 1; interorbital distance smaller than eye horizontal diameter; inner and outer metacarpal tubercle present).
The new species differs from R. dulongensis by (1) head wider than long; (2) interorbital distance larger than eye horizontal diameter; (3) nuptial pad present; (4) yellow disc; and (5) inner and outer metacarpal tubercle indistinct (vs. head smaller than long; interorbital distance smaller than eye horizontal diameter; nuptial pad absent; greyish or orange disc; inner and outer metacarpal tubercle indistinct present).
The new species differs from R. hillisi by (1) larger female body size (females 18.3–19.3 mm, n = 4); (2) head wider than long; (3) interorbital distance larger than eye horizontal diameter; (4) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V) ; and (5) and relative toe lengths: I < II < V < III < IV (vs. female 17.5 mm, n = 1; head longer than wider; interorbital distance smaller than eye horizontal diameter; webbing formula (II 1–2 III 1–21/2 IV 21/2–1 V); relative toe lengths: I < II < III < V <IV).
The new species differs from R. huanglianshan by (1) smaller female body size (females18.3–19.3 mm, n = 4); (2) lateral dermal fringe present; (3) yellow disc; (4) webbing formula (II 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); and (5) relative toe lengths: I < II < V < III < IV (vs. female 21.5 mm, n = 1; lateral dermal fringe absent; orange disc; fingers and toes lacking lateral dermal fringe; webbing formula (II 1–2 III 1 – 2 – IV 2 – 1 V); relative toe lengths: I < II < III < V < IV).
The new species differs from R. cangyuanensis by (1) interorbital distance larger than eye horizontal diameter; (2) nuptial pad small and milky white; and (3) yellow discs (vs. interorbital distance smaller than eye horizontal diameter; reddish nuptial pad at the base of first finger; orange disc).
The new species differs from R. ghatei by (1) smaller body size (males 14.6–17.7 mm, n = 7; females18.3–19.3 mm, n = 4); (2) tympanum distinct (TD 1.1–1.6 mm, n = 11); (3) nuptial pad present; (4) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); and (5) relative toe lengths: I < II < V < III < IV (vs. males 19.1–25.5 mm, n = 9; females 15.4–29.8 mm, n = 13; tympanum indistinct; nuptial pad absent; webbing formula (I 2 – 2 II 2 – 2½ III 2– 3 IV 2½ – 2 V); relative toe lengths: I < II < V = III < IV).
The new species differs from R. rezakhani by (1) smaller male body size (males 14.6–17.7 mm, n = 7); (2) interorbital distance larger than eye horizontal diameter; (3) tympanum distinct (TD 1.1–1.6 mm, n = 11); (4) nuptial pad present; (5) lateral dermal fringe present; (6) yellow disc; (7) inner and outer metacarpal tubercle indistinct; (8) inner metatarsal tubercle round; and (9) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V) (vs. males 18.8–19.0 mm; interorbital distance smaller than eye horizontal diameter; tympanum indistinct; nuptial pad absent; lateral dermal fringe absent; reddish or whitish; inner and outer metacarpal tubercle absent; inner metatarsal tubercle absent; webbing formula (I2 – 2 II 1¾ – 2 III 1½ – 3 IV 2¾ – 2 V).
The new species differs from R. annandalii by (1) head wider than long; and (2) relative toe lengths: I < II < V < III < IV (vs. head longer than wide; relative toe lengths: I < II < V = III < IV).
The new species differs from R. bombayensis by (1) smaller body size (males 14.6–17.7 mm, n = 7; females 18.3–19.3 mm, n = 4); (2) tympanum distinct (TD 1.1–1.6 mm, n = 11); and (3) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V) (vs. 30 mm, n = 1; tympanum indistinct; 1/3 webbing between toes).
The new species differs from R. tuberohumerus by (1) tympanum distinct (TD 1.1–1.6 mm, n = 11); and (2) relative toe lengths: I < II < V < III < IV (vs. tympanum indistinct; relative toe lengths: I < II ≤ V < III < IV).
The new species differs from R. longchuanensis by (1) smaller male body size (males 14.6–17.7 mm, n = 7); (2) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); and (3) yellow disc (vs. males 21.4–23.9 mm, n = 5; 1/4 webbing between toes; reddish, orange, or whitish disc).
The new species differs from R. andersoni by (1) interorbital distance larger than eye horizontal diameter; (2) webbing formula (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V); (3) yellow disc; and (4) relative toe lengths: I < II < V < III < IV (vs. interorbital distance smaller than eye horizontal diameter; 1/3 webbing between toes; orange disc; relative toe lengths: I < II < III = V < IV).
Recently,
Morphological comparison between Raorchestes malipoensis sp. nov. and Kurixalus gryllus (Smith, 1924).
Character | Species | |
---|---|---|
Raorchestes malipoensis sp. nov. (n = 11) | Kurixalus gryllus (n = 3) | |
SVL | 14.6–19.3 mm | 25.0–27.0 mm, |
HL | 5.2–7.9 mm | 8.0–9.5 mm |
HW | 5.5–8.2 mm | 10.0–11.0 mm |
EHD | 2.1–2.8 mm | 3.0–3.5 mm |
SL | 1.8–2.9 mm | 4–4.5 mm |
HAL | 4.2–5.6 mm | 7.5–8.5 mm |
TBL | 7.5–9.2 mm | 12–13 mm |
TD | 1.1–1.6 mm n = 11 | tympanum indistinct |
Tubercles along forearm and foot | absent | present |
Web of toes | I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V | toes a little more than half webbed |
Web of fingers | no webbing between fingers | fingers free except for a rudiment of a web between the two outer fingers |
Metatarsal tubercle | inner metatarsal tubercle rounded, outer metatarsal tubercle absent | a small inner metatarsal tubercle |
Coloration | dorsal surface beige, with pale brown and dark brown spots, an individual having large black spots on its body surface | dorsal color with pale or dark brown, green, yellow, or grey, many individuals had a bright green patch on the snout, and patches of similar color on the knees and round the vent |
In recent years, many new species have been found along the border between China and Vietnam, such as Odorrana geminata (
We thank reviewers for their comments on the manuscript. This work was supported by grants from the National Natural Science Foundation of China (32060114), Guangxi Natural Science Foundation Project (2022GXNSFAA035526), Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education (ERESEP2022Z04), ang Guangxi Key Laboratory of Rare and Endangered Animal Ecology, Guangxi Normal University (19-A-01-06).