Nidirana chapaensis (Bourret, 1937), one additional anuran species for the amphibian fauna of China

Abstract

The China-Vietnam border region is a well-known biodiversity hotspot, harboring a striking diversity of species and endemism. However, the largest part of this region is so far understudied due to restrictions by political boundaries and difficulties in accessibility. Consequently, many species have only been reported from one side of the political boundary, despite the presence of continuous habitat on both sides. In this study, we present our discovery of the frog Nidirana chapaensis in southern Yunnan Province, China. This species was previously known only from Vietnam. All major morphometric characters of these Chinese specimens were found to match with the Vietnamese specimens. In addition, our mitochondrial phylogeny suggests that the Chinese population is monophyletic with respect to topotypic Nidirana chapaensis, with uncorrected pairwise distances of 0.2% at the COI gene fragment analyzed. Therefore, we report Nidirana chapaensis as a new member of amphibian fauna of China, describe the morphological variation of the Chinese population, and provide additional natural history data of this species based on our observation from China.

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Accepted by M. Vences: 21 Feb. 2019; published: 28 Mar. 2019
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Article
https://doi.org/10.11646/zootaxa.4571.4.10
http://zoobank.org/urn:lsid:zoobank.org:pub:DBB3DB58-DD2A-4F0B-A101-F091C1C6006D
Nidirana chapaensis (Bourret, 1937), one additional anuran species for the
amphibian fauna of China
ZHIYONG YUAN
1,6,7
, XIAOLONG LIU
1,6
, KAI WANG
2,3
, JIAN WANG
4
, JINMIN CHEN
3
,
JIEQIONG JIN
3
, PINGFAN WEI
1
, JIAJUN ZHOU
5
& JING CHE
3,7
1
Key Laboratory for Conserving Wildlife with Small Populations in Yunnan, Southwest Forestry University, Kunming 650224, Yunnan,
China
2
Sam Noble Oklahoma Museum of Natural History & Department of Biology, University of Oklahoma, Norman, Oklahoma, USA
3
State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming
650223, Yunnan, China
4
College of Life Science and Technology, Honghe University, Mengzi 661199, Yunnan, China
5
Imaging Biodiversity Expedition Institute, Beijing100107, Beijing, China
6
These authors contributed equally
7
Corresponding authors. E-mail: chej@mail.kiz.ac.cn; yuanzhiyongkiz@126.com
Abstract
The China-Vietnam border region is a well-known biodiversity hotspot, harboring a striking diversity of species and en-
demism. However, the largest part of this region is so far understudied due to restrictions by political boundaries and dif-
ficulties in accessibility. Consequently, many species have only been reported from one side of the political boundary,
despite the presence of continuous habitat on both sides. In this study, we present our discovery of the frog Nidirana cha-
paensis in southern Yunnan Province, China. This species was previously known only from Vietnam. All major morpho-
metric characters of these Chinese specimens were found to match with the Vietnamese specimens. In addition, our
mitochondrial phylogeny suggests that the Chinese population is monophyletic with respect to topotypic Nidirana cha-
paensis, with uncorrected pairwise distances of 0.2% at the COI gene fragment analyzed. Therefore, we report Nidirana
chapaensis as a new member of amphibian fauna of China, describe the morphological variation of the Chinese popula-
tion, and provide additional natural history data of this species based on our observation from China.
Key words: new records, frog, China-Vietnam border, Yunnan Province
Introduction
As a direct result of increasing attentions and efforts in herpetological researches in China for the past decade, the
Chinese amphibian fauna has grown dramatically, with new records and new species described almost every year
(Murphy 2016). However, many regions of the country, particularly along the southern borders in Tibet, Yunnan,
and Guangxi, have not been surveyed in detail. Among these is the subtropical region along the China-Vietnam
border in Wenshan Prefecture of south-eastern Yunnan Province.
Music Frogs of the genus Nidirana comprise eight recognized species, including Nidirana adenopleura
(Boulenger 1909), N. chapaensis (Bourret 1937), N. daunchina (Chang 1933), N. hainanensis (Fei et al. 2007), N.
lini (Chou 1999), N. nankunensis (Lyu et al. 2017), N. okinavana (Boettger 1895), and N. pleuraden (Boulenger
1904). Most of these taxa are found in southern China (Fei et al. 2009). Among the three species that have not been
recorded in China is N. chapaensis, which occurs in northern Vietnam, Laos and Thailand, close to the southern
border of Yunnan Province. Given the continuous habitat that stretches across political boundaries in this region,
and its under-surveyed nature, it was considered likely that N. chapaensis was also found in the territory of China.
During the herpetological surveys of south-eastern Yunnan Province in 2016, we collected three Nidirana
individuals from a hitherto unknown population in Wenshan Prefecture (Fig. 1). Toes of seventeen additional frogs
were sampled. Following detailed morphological comparisons and phylogenetic analyses, we here confirm that the

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newly discovered Nidirana population in Yunnan should be assigned to N. chapaensis. Therefore, we report N.
chapaensis as a new amphibian record of China, provide descriptions of additional specimens, and provide remarks
on the natural history of the species based on our field observations.
FIGURE 1. Distribution of Nidirana chapaensis near the China-Vietnam border. Red circle (Xichou, Wenshan, Yunnan,
China) represent the new distribution record of the species in China.
Material and methods
Field sampling. Field surveys were conducted at Xiaoqiaogou Nature Reserve (23.362 ºN, 104.839 ºE, 1312 m
asl.), Xichou, Wenshan, Yunnan Province, China in April, June and August of 2016. A total of three specimens
were collected (males: KIZ 028112, KIZ 028136-37). Toes of seventeen additional individuals were clipped and
collected before releasing the living frogs in the locality of capture. The three collected individuals were euthanized
using a chlorobutanol solution, and liver or muscle tissues were taken from the specimens before fixing them in
75% ethanol. All specimens were deposited at the Museum of the Kunming Institute of Zoology (KIZ), Chinese
Academy of Sciences, Kunming, China.
Morphology and morphometrics. Measurements were taken of the three preserved individuals (KIZ028112,
KIZ028136-37) with a digital calliper to the nearest 0.1 mm by Zhiyong Yuan. Morphological characteristics and
measuring methods follow Fei et al. (2007). Abbreviations used are as follows: SVL (snout-vent length); HL(head
length); HW (head width); SNL (snout length); IND (internarial distance); IOD (interorbital distance at narrowest
point); EYE (diameter of the exposed portion of the eyeball); UEW (width of upper eyelid);TMP (horizontal
diameter of tympanum); TEY (tympanum–eye distance); HND (hand length); LAD (diameter of lower arm); FEM
(femur length); TIB (tibia length); TW (tibia width); TFL (length of tarsus and foot); FTL (foot length).
For comparative material, morphological measurements were obtained from recent studies of N. chapaensis,
which frogs from the newly discovered locality resembled (Luu et al. 2013; Ziegler et al. 2014). In addition,
morphological data of congeners was obtained from the literatures as well (Fei et al. 2007; Le et al. 2014; Iwai &
Shoda-Kagaya 2012; Fei et al. 2007).
DNA sequencing and analysis of sequences. To build a phylogeny for the genus Nidirana, we started by
extracting DNA from ethanol-preserved tissues of two individuals originating from the newly discovered locality
in Yunnan Province, and two individuals of N. hainanensis from Hainan Province (Table 1) using standard phenol-
chloroform extraction (Sambrook et al. 1989). A fragment of the mitochondrial gene for cytochrome oxidase
subunit 1 (COI) was amplified by polymerase chain reaction (PCRs). The primers and PCR conditions followed
Che et al. (2012). Then, we downloaded twelve available COI sequences from GenBank for the all the currently
recognized species of Nidirana excepted N. nankunensis, which was described recently and proved to sister with
the clade including N. adenopleura and N. okinavana (Lyu et al. 2017). Odorrana graminea was used as outgroup
based on a current phylogenetic hypothesis (Che et al. 2007). Sequences were checked for ambiguities and aligned
with the default settings (Edgar 2004). The interspecific and within-species mean uncorrected pairwise distances
were calculated in MEGA v6.0.6 (Tamura et al. 2013).

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TABLE 1 . Specimens used in the study, including their voucher numbers, localities, and GenBank accession numbers
for the COI gene.
We reconstructed the phylogeny of the genus Nidirana using both Maximum Likelihood (ML) and Bayesian
Inference (BI) analyses based on the COI sequences. The best-fit partitioning scheme and the best substitution
models for the respective partitions were identified with PartitionFinder v.1.1.1 (Lanfear et al. 2012) using the
Bayesian Information Criterion. A GTR + G model for all the three codon positions was selected as the best mode
and partition which was used in the ML and BI analyses. Maximum Likelihood analysis was conducted with
RAxML version 8.2.4 (Stamatakis 2014). The robustness of nodes of the best ML tree was assessed with 1000
bootstrap replicates. Bayesian Inference was executed in MrBayes v. 3.2.2 (Ronquist et al. 2012). The partitioned
analysis was conducted with two independent runs, each with four Metropolis coupled Markov Chain Monte Carlo
chains of 10 million generations and a burn-in of 25%. Convergence of sampled parameters (effective sampling
size/ESS for all parameters >200) was assessed by Tracer 1.6 (Rambaut et al. 2014).
Results and discussion
Morphological measurements of all the specimens from Yunnan are summarized in Table 2. All of the major
diagnostic characters of the Chinese specimens (KIZ 028112, KIZ 028136-37) match with the data presented in the
original description of N. chapaensis. Measurements of a few additional characters slightly differed from the
Vietnamese specimens (Luu et al. 2013; Ziegler et al. 2014), but nevertheless fell within close range of the latter
(Table 2).
A total of 555 base-pairs of new COI sequences were obtained from four samples obtained during field works
in Yunnan and Hainan (GenBank numbers MK492282–MK492285, Table 1). For all the alignment sequences, 187
variable positions and 163 parsimony informative sites were identified. Interspecific uncorrected pairwise
distances between the new population from Yunnan, China and the other species of Nidirana varied from 4.3%
(between N. hainanensis) to 11.5% (between N. pleuraden) (Table 3). The uncorrected pairwise genetic distance
between the new populations of Nidirana from Yunnan and the topotypic N. chapaensis (Sapa, Lao Cai, Vietnam)
was very small (0.9%). Furthermore, both the ML and BI analyses showed with strong support that the Yunnan
population forms a monophyletic clade with topotypic N. chapaensis (Fig. 2). Thus, based on evidence from both
molecular and morphological data, we conclude that the Nidirana population from Wenshan, Yunnan, China is
conspecific with N. chapaensis, and we report N. chapaensis as a new amphibian species occurring in China.
Species Voucher Locality GenBank Acc. No. Source
Nidirana adenopleura A-A-WZ001 unknown NC018771 Yu et al. 2012
Nidirana chapaensis 2000.485 Sapa, Lao Cai, Viet Nam KR087625 Grosjean et al. 2015
Nidirana chapaensis KIZ028136 Wenshan, Yunnan, China MK492282 This study
Nidirana chapaensis KIZ028137 Wenshan, Yunnan, China MK492283 This study
Nidirana daunchina SYNU12050568 Hangzhou, Zhejiang, China KF020615 Zhou et al. 2015
Nidirana hainanensis Tissue ID:
YPX19468
Diaoluoshan, Hainan, China MK492284 This study
Nidirana hainanensis Tissue ID:
YPX19469
Diaoluoshan, Hainan, China MK492285 This study
Nidirana lini unknown unknown HQ395352 Li et al. 2015
Nidirana okinavana unknown Ryukyu islands, Japan NC022872 Kakehashi et al. 2013
Nidirana pleuraden unknown Longling, Yunnan, China HQ395340 Li et al. 2015
Babina holsti unknown Ryukyu islands, Japan NC022870 Kakehashi et al. 2013
Babina subaspera AB679645 Amami islands, Japan AB679645 Iwai and Shodakagaya 2012
Babina subaspera AB761265 Amami islands, Japan AB761265 Iwai and Shodakagaya 2012
Odorrana graminea 2005.021 Long Nai Khao, Phongsali, Laos KR087839 Grosjean et al. 2015

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TABLE 2. Morphometric comparisons of Nidirana chapaensis populations from Yunnan, China (this study) and
Vietnam (Luu et al. 2013; Ziegler et al. 2014). Abbreviations are provided in the materials and method section. All
measurements are in the unit of millimeter (mm). “-” indicates without data.
TABLE 3. The pairwise uncorrected p-distance (%) of COI gene sequences between species of Nidirana. “-” indicates
genetic distance less than 0.1%.
Taxonomic conclusion
Nidirana chapaensis (Bourret, 1937)
(Figs. 3 and 4)
Description. Males: SVL 43.0–47.3mm; head longer than width (HL14.9–15.4 mm, HW 14.2–14.9 mm); snout
obtusely pointed in dorsal view, projecting beyond lower jaw; canthus rostralis distinct; nostril dorsolaterally
oriented, close to tip of snout; loreal region concave, sloping; tympanum distinct, rounded, smaller than eye
diameter (TMP = 58.2–69.6% EYE); supratympanic fold absent; pineal body distinct; internarial distance
relatively greater than interorbital distance (IOD = 86.4–96.5% IND). Two short rows of vomerine teeth present;
tongue notched posteriorly; a pair of internal vocal sacs present in males. Skin smooth, except small granular
tubercles present on posterior dorsum, near cloacae and hindlimbs; rictal glands distinct; dorsolateral fold distinct.
Catalog No. KIZ028112 KIZ028136 KIZ028137 Males from Quang Binh, Vietnam Male from Ha Giang, Vietnam
(IEBR A.2013.98)
SVL 47.3 46.6 43.0 43.0–46.0 42.1
HL 15.4 15.0 14.9 17.3–20.4 19.7
HW 14.9 14.4 14.2 14.9–18.0 15.2
SNL 7.4 7.5 6.6. 6.9–7.8. 6.9.
IND 5.8 5.9 5.7 5.1–6.3 5.1–6.3
IOD 5.4 5.1 5.5 3.3–4.5 3.3–4.5
UEW 3.3 4.2 3.3 - -
EYE 6.0 5.5 5.6 4.7–5.6 5.7
TMP 3.6 3.2 3.9 3.9–4.2 4.6
TEY 1.1 1.0 1.0 0.3–1.4 -
HND 18.2 18.9 15.9 - -
LAD 3.6 3.8 4.5 - -
FEM 18.3 22.9 19.4 21.4–25.2 -
TIB 21.0 23.8 21.0 24.5–29.5 -
TW 6.9 6.9 6.3 - -
TFL 34.0 35.8 32.2 - -
FTL 24.7 26.5 21.2 24.1–29.2 -
1234567
1N. adenopleura -
2N. chapaensis 0.096 -
3N. daunchina 0.047 0.099 -
4N. hainanensis 0.096 0.043 0.095 -
5N. lini 0.099 0.093 0.101 0.101 -
6N. okinavana 0.045 0.094 0.048 0.096 0.101 -
7N. pleuraden 0.106 0.115 0.111 0.122 0.111 0.110 -

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Relative finger length II < I <IV<III; fingers webbing absent; lateral fringes absent; finger tips expanded with
circummarginal grooves without discs; subarticular tubercles distinct; supernumerary tubercle indistinct; three oval
palmar tubercles present. Nuptial pad absent in males. Hind limbs long, tibia longer than thigh (TIB =87.1–96.2%
TFL); toes long and thin, relative lengths I<II<III<V<IV (Fig. 4); webbing present on toes, webbing formula: I 1–2
II 1–2 III 1–2 IV 2–1 V; narrowly lateral fringes distinct on toes; subarticular tubercles distinct; oval inner
metatarsal tubercle distinct; outer metatarsal tubercle indistinct; supernumerary tubercles absent on toes.
Coloration. In life, dorsal coloration of the adults ranges from orange to dark brown, with a narrow white
stripe running from pine body to vent, and a pattern of irregular black spots covering the dorsum. The loreal region
is blackish, and rictal glands are white. The region around tympanum is usually black, but can be creamy yellow
instead. The upper flank is blackish, with spots close to the dorsolateral fold. The remaining parts of flank are
mostly creamy yellow, sometimes with black spots scattered across. The iris is bronze. Dark bars are present on the
dorsal surface of the limbs. Ventral throat and chest are gray, covered densely with black spots; the belly is
yellowish or white. Ventral surfaces of the limbs are creamy-yellow or white with gray spots. The fingers have dark
gray spots; subarticular tubercles on the fingers and toes are pale gray and dark brown respectively, and the inner
metatarsal tubercle is creamy-yellow.
Distribution. The currently known ranges include northern Vietnam, Laos, Thailand, and south-eastern
Yunnan Province of China (Nguyen et al. 2009; Stuart 2005).
Natural history notes. Specimens were collected in small ponds adjacent to a stream (Fig. 5) between 19:00
and 23:30. The surrounding habitat was coniferous mixed forest at an elevation of ca. 1300 m a.s.l. Call of males
were heard at an air temperature of 24°C–28°C and a relative humidity of 53%–85% during April, June and
August, 2016. Many individuals were fund in April, but very few were found in June and August. Other frog
species observed at the same locality included Microhyla heymonsi, Limnonectes bannaensis, Fejervarya
multistriata, Sylvirana maosonensis.
FIGURE 2. Bayesian phylogenetic tree of the genus Nidirana inferred from a fragment of COI gene. “-” denotes low support
by Bayesian posterior probabilities (BPP < 95%), and bootstrap support (BS < 70%). The scale bar represents 0.05 nucleotide
substitutions per site.
On the distribution of herpetofauna across the China-Vietnam border. Our new record presented here
adds to the list of amphibian species which have recently been found to occur on both sides of the China-Vietnam
border, such as Limnonectes bannaensis (Suwannapoom et al. 2016), Paramesotriton deloustali (Zhang et al.
2017), and Tylototriton ziegleri (Jiang et al. 2017). Given that the area surrounding the China-Vietnam border is
located in the same zoogeographic region (Holt et al. 2013), and since habitats on both sides of the border are

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continuous without any known biogeographic barriers, it is highly possible that species many previously believed
endemic to either side of the border are found on the other side, and their distributions are much wider than
previously believed. Future collaborative research between herpetologists from both countries are needed in order
to obtain a better understanding of the biodiversity in this region.
FIGURE 3. Color variations of Nidirana chapaensis from the Yunnan population in China: (A–D) dorsolateral, dorsal (E), and
ventral (F) views of Nidirana chapaensis in life from China.
Acknowledgements
We thank Wouter Beukema, Miguel Vences and two anonymous reviewers for their help on English improve and
valuable suggestions during manuscript review. We thank Shaobin Hou, Baolin Zhang, Boyan Li, Jiansong Zhang
and the staff from Xiaoqiaogou nature reserve for helping with sample collections. This work was supported by
grants from the National Natural Science Foundation of China (31702008), Yunnan Applied Basic Research
Project (2018FD047), the First Class Discipline Construction Project for Forestry in Yunnan (No.51600625), and
the Biodiversity Conservation Program of Ministry of Environmental Protection of China to ZYY; Southeast Asia
Biodiversity Research Institute, CAS (Y4ZK111B01: 2017CASSEABRIQG002), China's Biodiversity
Observation Network (Sino-BON), and the Animal Branch of the Germplasm Bank of Wild Species, Chinese
Academy of Sciences (Large Research Infrastructure Funding) to JC.

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FIGURE 4. (A) Dorsal and (B) ventral views of the hand, and (C) dorsal and (D) ventral views of the foot of Nidirana
chapaensis in life from China.
FIGURE 5. (A) Habitat of Nidirana chapaensis in Xichou, Wenshan Prefecture, Yunnan Province, China.
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