Corresponding authors: Nikolay A. Poyarkov (
Academic editor: A. Crottini
Asian Mountain Toads (
Poyarkov Jr NA, Duong TV, Orlov NL, Gogoleva SS, Vassilieva AB, Nguyen LT, Nguyen VDH, Nguyen SN, Che J, Mahony S (2017) Molecular, morphological and acoustic assessment of the genus
Asian Mountain toads (
The systematic status of
The systematic status of the genus
Though a comprehensive phylogeny of the genus
For a long time after its’ description, the genus
The Langbian (or Da Lat) Plateau forms the southernmost edge of the Annamite Mountains, or Truong Son Range, a mountain chain spanning the breadth of Indochina, including parts of Vietnam, Laos and Cambodia. To date, following the review by
Distribution of
Measurements to the nearest 0.1 mm were taken using either a digital caliper, or a dissecting microscope; morphometrics of adult frogs and character terminology follows
Additionally, for the description of the type series we measured the distance between anterior orbital borders (
Morphological description of larval stages included the following 15 measurements: total length (
All statistical analyses were performed with Statistica 6.0 (StatSoft, Inc. 2001). Morphometric characters were used for univariate analyses, corrected by body size. Sexes were separated for subsequent comparisons among the samples. One-way ANOVA and Duncan’s post hoc test were used for morphometric comparisons. Multivariate statistical analyses were conducted for examination of overall morphological variation among studied populations. If some characters showed high correlation between each other, all but one of them were omitted in order to exclude the overweighting effect of these characters on the analyses. After metric values were log e-transformed, a principal component analysis (
Comparative morphological data were obtained from museum specimens of
We amplified sequences of a continuous fragment including partial sequences of 12S rRNA and 16S rRNA genes and complete t-val gene sequence, to obtain a fragment of up to 2077 bp (base pairs) of mtDNA. For some adult specimens and larvae a partial ca. 460–500 bp fragment of the 16S rRNA gene was sequenced for molecular identification purposes. 16S rRNA is a molecular marker widely applied for biodiversity surveys in amphibians (
Specimens and GenBank sequences of
GenBank |
Voucher ID | Species | Locality | Elevation (m a.s.l.) | Reference |
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500 |
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400 | this paper | ||
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400 | this paper | ||
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400 | this paper | ||
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400 | this paper | ||
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200 | this paper | ||
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830 | this paper | ||
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830 | this paper | ||
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1500 | this paper | ||
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1500 | this paper | ||
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1500 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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KIZ-013663** |
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1800 | this paper | |
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KIZ-013664** |
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1800 | this paper | |
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KIZ-013662** |
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1800 | this paper | |
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1500 | this paper | ||
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1500 | this paper | ||
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1500 | this paper | ||
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1500 | this paper | ||
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1500 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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1000 | this paper | ||
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2000 | this paper | |||
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2000 | this paper | |||
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2000 | this paper | |||
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2000 | this paper | |||
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2000 | this paper | |||
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2000 | this paper | |||
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2000 | this paper | |||
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1500 | this paper | |||
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KIZ YPX-05429 | 780 | this paper | ||
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KIZ YPX-05457 | 780 | this paper | ||
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KIZ YPX-05428 | 780 | this paper | ||
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1900 | this paper | |||
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1900 | this paper | |||
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2000 | this paper | |||
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1800 | this paper | |||
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1975 | this paper | |||
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DVT-00393 | 700 | this paper | ||
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AMNH A163680 |
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Vietnam, Quang Nam Prov., Tra My, Tra Don | 930 |
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AMNH A163669 |
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Vietnam, Quang Nam Prov., Tra My, Tra Don | 970 |
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Vietnam, Kon Tum Prov., Thac Nham forest | 1100 | this paper | |
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Vietnam, Gia Lai Prov., Kon Chu Rang N.R. | 1000 | this paper | |
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Vietnam, Gia Lai Prov., Kon Chu Rang N.R. | 1000 | this paper | |
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Vietnam, Gia Lai Prov., Kon Chu Rang N.R. | 1000 | this paper | |
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Vietnam, Gia Lai Prov., Kon Chu Rang N.R. | 1000 | this paper | |
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KUH 311601 |
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China, Guangxi Prov., Shiwan Dashang N.R., Fulong | 500 |
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AMNH A168682 |
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Vietnam, Lao Cai Prov., Van Ban Dist., Nam Tha | 330 |
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AMNH A163668 |
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Vietnam, Quang Nam Prov., Tra My, Tra Don | 980 |
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AMNH |
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Vietnam, Thua Thien–Hue Prov., A Luoi Dist., A Roang | 680 |
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ZP-AM 44 |
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– | – |
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Vietnam, Vinh Phuc Prov., Tam Dao | – | this paper | |
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Thailand, Satun Prov. | – | this paper | |
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Vietnam, Dien Bien Prov., Muong Nhe, Muong Nhe N.R. | – | this paper | |
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Thailand, Suratthani Prov. | – | this paper | |
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Vietnam, Dien Bien Prov., Muong Nhe, Muong Nhe N.R. | – | this paper | |
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ZYC1500 |
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China | – |
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ZYC1513 |
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China, Sichuan | – |
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DVT-04135 | Vietnam, Lao Cai Prov. | – | this paper | |
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Malaysia, Sarawak | – | this paper | |
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SCUM120630 |
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China, Sichuan Prov., Emei Shan Mt. | – |
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DVT-00298 |
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Vietnam, Phu Yen Prov., Tay Hoa | – | this paper |
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MVZ-Herp-223642 |
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– | – |
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Maximum Likelihood (
We
All recordings were standardized by Avisoft SASLab Pro software v. 5.2.05 in mono format with sampling rate at 48 kHz and 16-bit precision, and low-frequency noise was reduced using the low-pass filter (up to 1000 Hz). Calls were analyzed using Avisoft SASLab Pro software v. 5.2.05; all parameters were measured using the reticule and standard cursors in the spectrogram window of Avisoft. Spectrograms for analyses were created using the Hamming window, FFT-length 1024 points, frame 100%, and overlap 87.5%. Figure spectrograms were created using the Hamming window, FFT-length 512 points, frame 100%, and overlap 75%. In total, we measured 1797 calls of the new
Four temporal parameters were measured: the duration of each call, the interval between successive calls within each series, the duration of series, the interval between successive series, and five frequency parameters: the initial and final fundamental frequency, the minimum and maximum of fundamental frequency and the frequency of maximum amplitude (also “F peak”). Then we calculated the frequency range as the difference between the maximum and minimum of fundamental frequencies and the call repetition rate per recording/series (calls/s) for each recording/series as a ratio of number of all calls within the recording/series (excluding series consisting of just one call) to recording/series duration. All numerical parameters are given as mean ± SE, the minimum and maximum values are given in parentheses (min–max).
To compare acoustic characteristics between three species of
The records of advertisement calls were deposited at the Fonoteca Zoologica and are available at the website
The final alignment of the studied 12S rRNA–16S rRNA mtDNA gene fragment consisted of 2077 sites: 1439 sites were conserved and 567 sites were variable, of which 465 were found to be parsimony-informative. The transition–transversion bias (R) was estimated as 2.06. Nucleotide frequencies were A = 32.8%, T = 27.6%, C = 21.6%, and G = 17.9% (all data given for ingroup only).
We achieved high resolution of phylogenetic relationships among taxa within
Bayesian inference dendrogram of
Our analyses (Fig.
Our data confirm the monopyly of
Monophyly of the genus
Within Group I, the clade consisting of two specimens from central Vietnam (AMNH A-169287, Thua Thien-Hue Prov., and AMNH A-163668, Quang Nam Prov.: identified as
Within Group I, specimens of
Within Group II, Subclade A joins the medium-sized specimens from environs of the type locality of
The second species level clade comprises large-sized
The third species level clade forms a sister clade with respect to a clade comprised of
The observed interspecific sequence divergence within the genus
Uncorrected
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | |
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1.1 | 1.1 | 1.7 | 1.6 | 1.9 | 1.8 | 1.7 | 2.0 | 1.9 | 1.9 | 2.1 | 2.2 | 2.1 | 2.2 | 2.2 | 2.1 | 1.9 | 2.2 | 2.4 | |
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4.1 |
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0.9 | 1.6 | 1.6 | 1.9 | 1.8 | 1.8 | 1.8 | 1.8 | 1.7 | 2.0 | 2.1 | 2.1 | 2.2 | 2.2 | 2.0 | 1.9 | 2.2 | 2.3 | |
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5.0 | 2.6 |
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1.7 | 1.6 | 1.8 | 1.9 | 1.7 | 1.8 | 1.8 | 1.6 | 2.0 | 2.1 | 2.1 | 2.1 | 2.1 | 2.0 | 1.9 | 2.2 | 2.2 | |
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9.1 | 9.3 | 10.0 |
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0.9 | 1.8 | 1.6 | 1.7 | 2.1 | 1.8 | 1.9 | 2.2 | 2.0 | 2.2 | 2.2 | 2.3 | 2.1 | 2.1 | 2.3 | 2.2 | |
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8.2 | 8.9 | 8.7 | 3.1 |
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1.8 | 1.6 | 1.6 | 2.1 | 2.0 | 2.1 | 2.2 | 1.8 | 2.3 | 2.2 | 2.1 | 2.1 | 2.0 | 2.3 | 2.3 | |
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10.9 | 11.8 | 11.6 | 9.7 | 9.5 |
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1.3 | 1.5 | 1.6 | 1.9 | 1.7 | 1.7 | 1.8 | 1.9 | 2.1 | 2.2 | 2.0 | 1.9 | 2.4 | 2.4 |
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10.9 | 12.6 | 13.0 | 9.7 | 10.0 | 7.6 |
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1.3 | 1.7 | 1.8 | 1.7 | 1.9 | 1.8 | 2.1 | 2.2 | 2.3 | 1.9 | 1.8 | 2.4 | 2.5 |
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9.1 | 10.5 | 10.6 | 8.6 | 9.4 | 7.8 | 7.3 |
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1.8 | 1.7 | 1.7 | 1.8 | 1.7 | 2.0 | 2.1 | 2.5 | 1.9 | 1.7 | 2.2 | 2.2 |
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12.8 | 13.0 | 12.7 | 10.3 | 10.9 | 8.0 | 10.8 | 8.8 | – | 1.8 | 1.4 | 1.9 | 2.0 | 2.0 | 2.2 | 2.1 | 1.5 | 1.7 | 2.3 | 2.4 |
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11.6 | 12.2 | 11.9 | 11.1 | 11.7 | 10.3 | 9.5 | 8.8 | 7.8 | – | 1.3 | 1.8 | 1.8 | 1.8 | 2.2 | 2.4 | 1.7 | 1.7 | 2.2 | 2.2 | |
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10.8 | 11.4 | 11.0 | 9.9 | 10.2 | 8.8 | 9.6 | 8.4 | 5.3 | 3.7 | – | 1.7 | 1.8 | 1.8 | 2.1 | 2.1 | 1.2 | 1.6 | 2.2 | 2.2 |
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13.6 | 14.7 | 15.1 | 14.4 | 15.2 | 10.5 | 12.8 | 9.7 | 11.1 | 10.3 | 9.9 | – | 1.4 | 1.4 | 1.9 | 2.2 | 1.8 | 1.7 | 2.4 | 2.4 |
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13.6 | 14.2 | 14.3 | 11.1 | 10.7 | 10.1 | 11.4 | 9.7 | 9.5 | 9.1 | 9.1 | 7.4 | – | 1.6 | 1.9 | 2.3 | 1.7 | 1.8 | 2.5 | 2.4 |
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12.8 | 13.4 | 14.3 | 14.8 | 14.5 | 11.3 | 14.0 | 11.5 | 11.1 | 9.5 | 9.1 | 6.6 | 6.2 | – | 2.0 | 2.3 | 2.0 | 2.1 | 2.6 | 2.3 |
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16.5 | 16.3 | 16.0 | 18.1 | 18.3 | 16.9 | 18.9 | 14.8 | 14.8 | 14.0 | 12.8 | 13.6 | 14.4 | 13.6 | – | 2.5 | 2.0 | 1.9 | 2.3 | 2.4 |
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14.5 | 16.0 | 16.4 | 16.0 | 14.5 | 15.8 | 17.8 | 15.8 | 14.0 | 14.4 | 13.6 | 13.6 | 13.2 | 14.8 | 16.5 | – | 2.0 | 2.2 | 2.6 | 2.4 |
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12.8 | 13.0 | 13.7 | 10.7 | 11.5 | 11.5 | 11.7 | 10.9 | 7.0 | 7.4 | 5.3 | 11.1 | 9.1 | 11.1 | 12.3 | 14.4 | – | 1.4 | 2.4 | 2.4 |
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11.5 | 11.4 | 12.0 | 11.5 | 11.5 | 11.5 | 11.3 | 10.1 | 8.6 | 7.4 | 7.4 | 10.3 | 9.5 | 11.9 | 12.3 | 12.8 | 4.9 | – | 2.2 | 2.3 |
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17.5 | 18.8 | 19.4 | 18.5 | 19.9 | 21.0 | 21.1 | 18.9 | 21.8 | 19.8 | 21.0 | 20.2 | 19.3 | 21.4 | 22.6 | 23.9 | 20.2 | 18.9 | – | 2.2 |
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23.1 | 23.9 | 24.0 | 22.2 | 21.8 | 23.5 | 24.7 | 23.1 | 22.8 | 22.0 | 21.8 | 23.3 | 22.4 | 23.7 | 23.7 | 22.2 | 20.4 | 20.8 | 21.8 | – |
Intraspecific distances within
The newly discovered lineage of
Among the three species examined, mean
Boxplots of
Measurements of the three species of
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Males | 35.9 ± 3.4 | 10.3 ± 0.8 | 10.2 ± 0.9 | 4.4 ± 0.4 | 2.4 ± 0.4 | 2.0 ± 0.3 | 3.2 ± 0.5 | 1.4 ± 0.3 | 1.6 ± 0.3 | 3.3 ± 0.5 | 2.7 ± 0.4 | 3.1 ± 0.4 | 8.7 ± 1.0 | 8.7 ± 0.9 | 3.7 ± 0.6 | 3.7 ± 0.4 | 5.9 ± 0.6 | 3.5 ± 0.6 | 17.2 ± 2.8 | 16.8 ± 2.5 | 14.7 ± 2.4 | 23.7 ± 2.6 | 2.0 ± 0.5 |
(31.7–42.2) | (9.2–12.0) | (9.0–12.0) | (3.8–5.3) | (1.8–3.1) | (1.7–2.5) | (2.6–3.8) | (0.7–1.8) | (1.1–2.2) | (2.2–4.1) | (1.9–3.4) | (2.2–3.8) | (7.2–11.0) | (7.2–10.1) | (2.7–5.0) | (3.1–4.4) | (4.2–7.2) | (2.3–4.6) | (14.4–24.6) | (13.9–23.3) | (11.2–18.8) | (19.2–27.6) | (0.9–2.7) | |
Females | 45.1 ± 2.2 | 11.3 ± 0.4 | 11.4 ± 0.6 | 4.8 ± 0.4 | 2.7 ± 0.2 | 2.5 ± 0.2 | 3.4 ± 0.4 | 1.6 ± 0.3 | 1.7 ± 0.4 | 3.8 ± 0.2 | 2.8 ± 0.1 | 3.9 ± 0.3 | 10.4 ± 0.6 | 10.5 ± 1.4 | 4.7 ± 0.7 | 4.3 ± 0.6 | 7.2 ± 0.5 | 4.6 ± 0.7 | 19.7 ± 1.0 | 19.6 ± 1.6 | 16.7 ± 0.5 | 27.0 ± 0.9 | 2.9 ± 0.2 |
(43.1–47.4) | (10.9–11.7) | (10.9–12.1) | (4.4–5.1) | (2.5–3.0) | (2.3–2.6) | (2.9–3.6) | (1.2–1.9) | (1.4–2.2) | (3.6–4.0) | (2.7–2.9) | (3.5–4.2) | (9.8–10.9) | (9.0–11.9) | (3.9–5.3) | (3.8–4.9) | (6.7–7.7) | (4.1–5.4) | (18.7–20.7) | (17.8–20.6) | (16.3–17.2) | (26.0–27.9) | (2.6–3.0) | |
Males | 29.7 ± 1.8 | 8.6 ± 0.7 | 8.9 ± 0.6 | 3.7 ± 0.3 | 2.2 ± 0.2 | 1.5 ± 0.2 | 2.7 ± 0.4 | 1.2 ± 0.2 | 1.4 ± 0.3 | 2.7 ± 0.3 | 2.3 ± 0.5 | 2.9 ± 1.0 | 7.3 ± 0.8 | 7.0 ± 0.9 | 2.6 ± 0.5 | 2.9 ± 0.4 | 4.8 ± 0.5 | 3.0 ± 0.6 | 14.0 ± 1.7 | 13.7 ± 1.3 | 11.9 ± 1.0 | 19.8 ± 2.2 | 1.6 ± 0.4 |
(26.9–33.9) | (7.2–10.1) | (7.4–9.8) | (3.1–4.3) | (1.6–2.7) | (1.1–2.1) | (2.0–3.3) | (0.8–1.6) | (1.0–2.2) | (2.1–3.4) | (1.5–3.3) | (2.4–7.9) | (6.1–8.9) | (4.8–9.4) | (1.1–3.5) | (2.0–4.1) | (3.7–6.0) | (1.4–4.1) | (11.7–19.6) | (11.3–16.1) | (10.4–14.7) | (17.1–28.0) | (1.2–2.5) | |
Females | 35.6 ± 0.5 | 9.7 ± 0.5 | 9.8 ± 1.1 | 4.0 ± 0.2 | 2.3 ± 0.4 | 1.7 ± 0.1 | 3.1 ± 0.5 | 1.4 ± 0.3 | 1.7 ± 0.3 | 2.9 ± 0.5 | 2.5 ± 0.4 | 2.9 ± 0.2 | 9.0 ± 1.1 | 8.8 ± 0.7 | 3.7 ± 0.7 | 3.9 ± 0.5 | 5.8 ± 0.8 | 4.1 ± 0.9 | 17.1 ± 2.0 | 16.3 ± 1.0 | 14.9 ± 0.7 | 24.3 ± 2.1 | 2.0 ± 0.3 |
(35.1–36.5) | (9.0–10.2) | (8.6–11.0) | (3.6–4.2) | (1.9–3.0) | (1.6–1.9) | (2.4–3.6) | (1.0–1.7) | (1.2–2.1) | (2.3–3.6) | (1.7–2.9) | (2.6–3.1) | (7.7–10.1) | (7.7–9.7) | (2.9–4.5) | (3.3–4.7) | (4.9–7.0) | (3.1–5.2) | (14.9–19.8) | (15.5–17.8) | (13.9–15.7) | (21.8–27.2) | (1.5–2.4) | |
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Males | 45.7 ± 4.3 | 13.8 ± 0.9 | 13.5 ± 1.0 | 5.1 ± 0.3 | 3.5 ± 0.5 | 2.8 ± 0.4 | 3.8 ± 0.5 | 1.9 ± 0.4 | 1.6 ± 0.3 | 4.5 ± 0.6 | 3.1 ± 0.3 | 3.7 ± 0.4 | 12.6 ± 1.3 | 11.0 ± 1.1 | 4.6 ± 0.6 | 4.7 ± 0.6 | 7.5 ± 0.6 | 4.5 ± 0.6 | 20.6 ± 1.6 | 20.7 ± 2.2 | 18.4 ± 2.0 | 29.3 ± 2.8 | 2.5 ± 0.4 |
(38.2–53.7) | (12.5–15.6) | (12.2–15.4) | (4.7–5.6) | (3.0–4.7) | (1.9–3.6) | (3.0–4.6) | (1.3–2.5) | (1.2–2.2) | (3.5–5.5) | (2.5–3.5) | (3.0–4.1) | (10.4–14.7) | (9.2–11.1) | (3.7–5.8) | (3.7–5.6) | (6.2–8.4) | (3.6–5.5) | (17.7–23.3) | (18.1–25.1) | (15.0–22.4) | (24.0–33.5) | (1.9–3.1) | |
Females | 60.8 ± 9.7 | 16.6 ± 1.8 | 16.1 ± 1.8 | 5.3 ± 0.4 | 3.7 ± 0.2 | 3.3 ± 0.3 | 4.0 ± 0.3 | 2.0 ± 0.4 | 1.8 ± 0.4 | 4.7 ± 0.2 | 3.5 ± 0.6 | 4.1 ± 0.2 | 14.3 ± 1.0 | 13.7 ± 3.3 | 5.1 ± 0.5 | 5.9 ± 1.0 | 8.4 ± 0.9 | 5.1 ± 1.4 | 23.4 ± 2.0 | 24.3 ± 1.5 | 20.7 ± 4.6 | 32.3 ± 5.0 | 3.2 ± 0.4 |
(51.4–70.7) | (14.6–18.0) | (14.0–17.4) | (5.0–5.7) | (3.4–3.9) | (3.0–3.6) | (3.7–4.3) | (1.6–2.4) | (1.3–2.2) | (4.4–4.9) | (3.0–4.1) | (3.8–4.2) | (13.6–15.5) | (11.0–17.4) | (4.7–5.7) | (5.1–7.0) | (7.6–9.3) | (4.3–6.7) | (21.3–25.3) | (22.7–25.7) | (15.7–24.8) | (26.8–36.4) | (2.8–3.6) |
The results of the multivariate
Two-dimensional plots of
In summary, both in body size and other meristic characters, all three examined lineages of Langbian
Measurements of advertisement call parameters for three
Measurements of advertisement call parameters for three species of
Parameters |
|
Tukey/ |
|
Tukey/ |
ANOVA/ |
|
---|---|---|---|---|---|---|
Number of males | 3 | 3 | – | 3 | – | – |
Number of recordings | 5 | 3 | – | 3 | – | – |
Number of series | 140 | 115 | – | 15 | – | – |
Number of calls | 1797 | 533 | – | 200 | – | – |
Call repetition rate per recording (calls/ |
1.18 ± 0.2 |
0.35 ± 0.14 |
p < 0.05 | 3.07 ± 0.13 |
p < 0.001 | |
Number of calls per series | 12.84 ± 0.41 |
4.64 ± 0.16 |
p |
13.33 ± 1.4 |
p < 0.05 | |
Series duration ( |
3.42 ± 0.11 |
2.18 ± 0.09 |
p < 0.001 | 2.59 ± 0.33 |
p = 0.88 | |
Call repetition rate per series (calls/ |
3.87 ± 0.07 |
2.33 ± 0.03 |
p < 0.001 | 5.34 ± 0.15 |
p < 0.001 | |
Call duration ( |
73 ± 0.23 |
104 ± 0.56 |
p < 0.001 |
62 ± 0.46 |
p < 0.001 |
H2.2530 = 1345.1* |
Inter-calls interval ( |
207 ± 2.06 |
421.54 ± 4.17 |
p < 0.001 |
143 ± 3.32 |
p < 0.001 |
H2.2260 = 1008.5* |
Inter-series interval ( |
6.51 ± 0.41 |
7.98 ± 0.55 |
p < 0.001 |
1.64 ± 0.19 |
p < 0.001 |
H2.259 = 42.7* |
4348.02 ± 2.96 |
4414.17 ± 5.12 |
p < 0.001 |
3449.55 ± 6.41 |
p < 0.001 |
H2.2530 = 655.7* | |
4715.3 ± 3.29 |
4888.76 ± 3.82 |
p < 0.001 |
3708.9 ± 9.28 |
p < 0.001 |
H2.2530 = 1075.9* | |
4807.74 ± 3.46 |
4998.74 ± 4.27 |
p < 0.001 |
3907.05 ± 4.22 |
p < 0.001 |
H2.2530 = 1094.2* | |
4348.02 ± 2.96 |
4414.17 ± 5.12 |
p < 0.001 |
3449.55 ± 6.41 |
p < 0.001 |
H2.2530 = 689.5* | |
459.71 ± 3.27 |
584.58 ± 3.87 |
p < 0.001 |
457.5 ± 5.86 |
p = 0.61 |
H2.2530 = 367.1* | |
4645.94 ± 4.39 |
4845.99 ± 4.22 |
p < 0.001 |
3798.9 ± 4.87 |
p < 0.001 |
H2.2530 = 1030.2* |
Advertisement calls of all studied species were similarly uttered in series (Fig.
Oscillograms (top) and sonograms (bottom) of male advertisement calls of
The frequency of maximum amplitude always coincided with the fundamental frequency and greatly varied within recordings: from 4030 to 4920 Hz for the undescribed
Number of harmonics varied between/within recordings but this characteristic mostly depended on recording quality (e.g., sensitivity of recording equipment, distance from vocalizing animal, signal volume and background noise). Calls from the highest quality recordings (of the undescribed
We had two sets of the undescribed
Our study, based on three lines of evidence — phylogenetic analysis and distribution of mtDNA haplotypes (Figs
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English: “
Mature male (
Holotype
Head moderately small (
Forelimbs slender, forearm moderately long (
Hindlimbs slender, relatively long, shanks overlap when thighs are held at right angle to body; shank length less than half of snout to vent length (
Skin of dorsal and lateral surfaces of head, body and limbs smooth with numerous small tubercles finely and relatively evenly scattered on dorsal surfaces of trunk, head and limbs (Fig.
(Figure
“
[?]
The studied specimens of
English: “
Based upon several lines of evidence, including the analyses of diagnostic morphological characters, acoustic analyses of advertisement calls and phylogenetic analyses of mtDNA sequences for the 12S rRNA–16S rRNA genes, the new species of
“
[?]
“
Holotype
The specific epithet is an adjective (in agreement with the genus name in feminine gender), derived from “
The recommended common name in English is “Elfin Mountain Toad”. The recommended common name in Vietnamese is “Cóc Núi Tiểu Yêu Tinh”.
The species is allocated to
The new species is also markedly distinct from all congeners for which comparable sequences are available (16S rRNA mitochondrial gene; uncorrected genetic distance > 8.2%). The advertisement call of the new species consists of whistling notes uttered in series: average 12.84 ± 0.41 calls per series, with an average dominant frequency of 4645.94 ± 4.39 Hz, repetition rate per recording/series 1.18 ± 0.2 calls/s and 3.87 ± 0.07 calls/s, respectively, with average call duration 73 ± 0.23 ms and inter-call interval 207 ± 2.06 ms, also distinguishes the new species from
Mature male (
Holotype of
Head moderately large (
Forelimbs slender, forearm moderately long (
Hindlimbs slender, relatively long, shanks overlap when thighs are held at right angle to body; shank length less than half of snout–vent length (
Skin of dorsal and lateral surfaces of head, body and limbs shagreened, with numerous small skin asperities present on anterior two thirds of dorsum, sparse posteriorly, increasing in density along dermal ridges, densely covering dorsal and lateral surfaces of head, upper eyelids, and dorsal surfaces of thighs, shanks, upper forelimbs, forearms, hands, feet and digits, and absent from all remaining surfaces. Small tubercles finely and relatively evenly scattered on dorsal surfaces of trunk, head and limbs, including maxilla, mandible, eyelids and dorsal surfaces of head, forelimbs and hindlimbs (Figs
Entire dorsum light olive-brown to yellow-brown with large irregular brownish grey spots; dorsal surfaces of head yellowish brown from tip of snout to eyes; small oval-shaped spot with irregular borders on dorsal surface of snout between anterior canthi; a small dark dot on dorsal surface of snout tip; similar single dark dots on anterior parts of upper eyelids; brown “V”-shaped marking on crown between supraorbital horns with apex pointing posteriorly, outlined with thin light-beige edging; round brownish spot at head basis; “ >–< ”-shaped marking surrounded with dark olive-brown, outlined with thin light-beige edging forming a hourglass-shaped dorsal marking (Fig.
In preservative coloration faded to light grey-brown on dorsum and flanks, with slightly paler limbs and greyish beige to whitish on venter; reddish and orange tints, as well as iris coloration, faded completely; dark markings on dorsum, sides and venter and other features remain without significant change (Fig.
Morphometric variation within the type series and other referred specimens of the new species is shown in Table
Paratypes of
Measurements of the
Specimen ID | Population | Sex | Type status |
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NAP-02658 | 10 | m | holotype | 27.2 | 8.0 | 7.9 | 3.6 | 1.9 | 1.6 | 2.5 | 1.3 | 1.3 | 2.3 | 2.3 | 2.0 |
NAP-01455 | 6 | m | paratype | 33.9 | 10.1 | 9.8 | 3.8 | 2.3 | 1.9 | 2.7 | 1.3 | 1.3 | 2.8 | 3.1 | 2.9 |
NAP-01449 | 6 | m | paratype | 31.0 | 9.3 | 9.8 | 3.8 | 2.2 | 1.6 | 2.5 | 1.3 | 1.2 | 3.1 | 2.9 | 3.5 |
NAP-01450 | 6 | m | paratype | 29.2 | 9.0 | 9.0 | 3.7 | 2.3 | 1.4 | 3.0 | 1.2 | 1.4 | 3.2 | 2.5 | 3.2 |
NAP-01460 | 6 | m | paratype | 29.6 | 8.7 | 8.4 | 3.4 | 2.7 | 1.1 | 2.1 | 1.1 | 1.0 | 2.5 | 1.5 | 2.6 |
ABV-00316 | 8 | m | 28.9 | 8.8 | 9.1 | 3.8 | 2.6 | 1.6 | 2.2 | 1.3 | 1.1 | 2.8 | 1.8 | 3.1 | |
HB-36-1 | 8 | m | 27.5 | 8.6 | 9.2 | 4.1 | 2.4 | 1.3 | 2.6 | 1.4 | 1.4 | 2.6 | 2.1 | 2.6 | |
ROM-36525 | 11 | m | 26.9 | 7.6 | 8.1 | 3.6 | 1.9 | 1.3 | 2.9 | 1.3 | 1.2 | 2.3 | 1.7 | 2.8 | |
ROM-36523 | 11 | m | 27.8 | 8.2 | 8.4 | 4.0 | 2.2 | 1.8 | 2.8 | 0.9 | 1.5 | 2.1 | 1.6 | 2.9 | |
ROM-36529 | 11 | m | 28.3 | 7.9 | 8.6 | 4.3 | 2.2 | 1.6 | 2.4 | 1.1 | 1.5 | 2.8 | 2.0 | 2.7 | |
NAP-01757 | 6 | m | 32.0 | 9.2 | 9.2 | 4.0 | 2.3 | 1.8 | 2.6 | 1.3 | 1.3 | 2.3 | 2.3 | 2.5 | |
NAP-01782 | 6 | m | 31.1 | 9.3 | 9.1 | 3.8 | 2.2 | 1.7 | 2.3 | 1.1 | 1.0 | 2.8 | 2.6 | 7.9 | |
NAP-01758 | 7 | m | 32.1 | 9.2 | 9.3 | 3.4 | 2.4 | 1.7 | 2.4 | 0.8 | 1.3 | 3.1 | 2.0 | 2.4 | |
NAP-01871 | 7 | m | 32.8 | 8.8 | 9.6 | 3.7 | 2.2 | 1.3 | 2.7 | 0.9 | 1.1 | 2.9 | 1.6 | 2.4 | |
NAP-01783 | 7 | m | 31.3 | 9.0 | 9.2 | 4.0 | 2.2 | 1.6 | 2.8 | 1.2 | 1.1 | 3.1 | 2.1 | 2.8 | |
ABV-00471 | 6 | m | paratype | 30.7 | 8.5 | 8.5 | 3.7 | 2.3 | 1.3 | 2.6 | 0.9 | 1.4 | 2.7 | 2.4 | 2.7 |
ABV-00454 | 6 | m | paratype | 31.4 | 8.8 | 9.3 | 3.8 | 2.1 | 1.5 | 2.0 | 1.0 | 1.1 | 2.5 | 1.9 | 2.9 |
ABV-00472 | 6 | m | paratype | 30.8 | 8.9 | 9.3 | 3.3 | 2.3 | 2.1 | 2.3 | 1.6 | 1.0 | 3.2 | 2.1 | 2.6 |
ROM-36524 | 11 | m | 28.5 | 8.3 | 8.5 | 3.6 | 2.1 | 1.2 | 2.8 | 0.8 | 1.6 | 2.4 | 2.2 | 2.5 | |
ROM-36527 | 11 | m | 27.7 | 7.4 | 7.8 | 3.1 | 2.0 | 1.7 | 3.0 | 1.3 | 1.8 | 2.7 | 2.1 | 2.7 | |
ROM-36522 | 11 | m | 28.1 | 7.9 | 8.4 | 3.2 | 1.7 | 1.3 | 2.8 | 1.2 | 1.5 | 2.6 | 2.1 | 2.8 | |
ROM-36528 | 11 | m | 28.2 | 7.2 | 7.4 | 3.1 | 1.6 | 1.5 | 3.3 | 1.0 | 1.8 | 3.4 | 1.9 | 2.9 | |
ROM-36526 | 11 | m | 27.1 | 7.6 | 8.2 | 3.6 | 2.1 | 1.5 | 2.9 | 1.1 | 2.1 | 2.6 | 2.1 | 2.4 | |
8 | m | 27.9 | 8.2 | 8.9 | 3.9 | 2.2 | 1.2 | 2.5 | 1.1 | 1.4 | 2.1 | 2.8 | 3.1 | ||
8 | m | 29.4 | 9.6 | 9.1 | 3.4 | 2.3 | 1.4 | 3.3 | 1.1 | 2.2 | 2.7 | 3.3 | 2.7 | ||
8 | m | 30.1 | 9.5 | 9.8 | 4.1 | 2.2 | 1.8 | 2.9 | 1.3 | 1.6 | 2.4 | 3.3 | 2.7 | ||
8 | m | 29.6 | 8.7 | 9.3 | 3.4 | 2.2 | 1.3 | 2.8 | 1.1 | 1.7 | 2.2 | 2.9 | 2.7 | ||
8 | m | 30.8 | 9.2 | 9.6 | 3.4 | 2.1 | 1.6 | 3.3 | 1.4 | 1.9 | 2.4 | 2.6 | 2.6 | ||
8 | m | 28.6 | 8.7 | 8.9 | 3.5 | 2.3 | 1.8 | 3.3 | 1.2 | 2.1 | 2.6 | 2.8 | 2.5 | ||
ABV-00455 | 6 | f | paratype | 35.2 | 9.3 | 8.9 | 4.2 | 2.4 | 1.7 | 2.4 | 1.7 | 1.2 | 2.8 | 1.7 | 2.6 |
CYS-10-10 | 11 | f | 35.1 | 9.4 | 9.1 | 3.9 | 1.9 | 1.9 | 3.6 | 1.6 | 2.1 | 3.5 | 2.6 | 3.0 | |
ROM-36530 | 11 | f | 35.7 | 9.0 | 8.6 | 3.6 | 2.1 | 1.6 | 3.3 | 1.0 | 1.9 | 3.6 | 2.4 | 2.8 | |
8 | f | 36.5 | 10.0 | 11.0 | 4.1 | 2.4 | 1.8 | 2.7 | 1.2 | 1.5 | 2.7 | 2.8 | 3.1 | ||
8 | f | 35.8 | 10.1 | 10.9 | 4.2 | 3.0 | 1.6 | 3.2 | 1.3 | 1.9 | 2.3 | 2.6 | 2.8 | ||
8 | f | 35.3 | 10.2 | 10.3 | 4.1 | 2.2 | 1.8 | 3.3 | 1.5 | 1.8 | 2.8 | 2.9 | 3.1 | ||
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NAP-02658 | 10 | m | holotype | 7.0 | 7.3 | 2.4 | 2.8 | 4.6 | 2.8 | 13.2 | 12.1 | 18.9 | 11.9 | 1.8 | |
NAP-01455 | 6 | m | paratype | 8.5 | 9.4 | 2.3 | 4.1 | 6.0 | 3.7 | 16.1 | 14.8 | 14.7 | 28.0 | 2.0 | |
NAP-01449 | 6 | m | paratype | 7.1 | 7.5 | 2.8 | 3.0 | 4.1 | 3.0 | 14.4 | 13.4 | 11.8 | 19.4 | 1.7 | |
NAP-01450 | 6 | m | paratype | 7.5 | 7.8 | 2.1 | 2.9 | 4.9 | 2.6 | 12.9 | 13.8 | 11.1 | 19.7 | 1.9 | |
NAP-01460 | 6 | m | paratype | 6.5 | 6.7 | 2.3 | 2.9 | 4.4 | 2.4 | 13.4 | 11.3 | 10.6 | 17.5 | 1.3 | |
ABV-00316 | 8 | m | 7.3 | 6.7 | 2.4 | 2.9 | 4.9 | 2.9 | 13.2 | 13.5 | 11.6 | 19.2 | 2.5 | ||
HB-36-1 | 8 | m | 6.6 | 7.1 | 2.1 | 3.2 | 4.6 | 3.0 | 12.6 | 13.4 | 11.6 | 18.9 | 1.5 | ||
ROM-36525 | 11 | m | 6.9 | 6.6 | 2.0 | 2.4 | 4.5 | 2.7 | 11.7 | 11.9 | 10.6 | 17.1 | 1.2 | ||
ROM-36523 | 11 | m | 6.4 | 6.0 | 2.9 | 2.9 | 4.8 | 3.0 | 12.8 | 13.2 | 11.4 | 18.3 | 1.6 | ||
ROM-36529 | 11 | m | 6.7 | 6.4 | 2.9 | 3.0 | 4.5 | 3.0 | 13.5 | 13.5 | 11.0 | 19.1 | 1.6 | ||
NAP-01757 | 6 | m | 8.4 | 7.4 | 2.5 | 2.5 | 4.5 | 3.1 | 13.3 | 14.5 | 11.6 | 19.6 | 1.2 | ||
NAP-01782 | 6 | m | 6.1 | 4.8 | 1.1 | 2.0 | 3.7 | 1.4 | 12.7 | 13.1 | 11.0 | 18.7 | 1.4 | ||
NAP-01758 | 7 | m | 8.1 | 6.6 | 2.1 | 3.2 | 4.6 | 2.5 | 14.8 | 15.3 | 13.3 | 20.4 | 1.8 | ||
NAP-01871 | 7 | m | 6.5 | 6.8 | 2.3 | 2.8 | 4.6 | 3.1 | 14.2 | 14.6 | 10.8 | 19.2 | 1.2 | ||
NAP-01783 | 7 | m | 7.6 | 6.1 | 2.4 | 3.4 | 4.8 | 3.0 | 14.0 | 15.7 | 10.4 | 18.1 | 1.6 | ||
ABV-00471 | 6 | m | paratype | 8.6 | 7.1 | 3.0 | 3.4 | 4.9 | 3.2 | 14.2 | 14.5 | 13.1 | 19.6 | 1.2 | |
ABV-00454 | 6 | m | paratype | 8.2 | 7.2 | 3.0 | 3.2 | 5.1 | 3.2 | 15.7 | 15.3 | 13.1 | 22.0 | 1.3 | |
ABV-00472 | 6 | m | paratype | 7.9 | 6.5 | 2.5 | 3.0 | 5.2 | 3.1 | 14.7 | 11.9 | 12.6 | 21.4 | 1.2 | |
ROM-36524 | 11 | m | 6.8 | 6.4 | 2.2 | 2.9 | 4.5 | 2.4 | 13.0 | 13.0 | 10.8 | 17.4 | 1.5 | ||
ROM-36527 | 11 | m | 6.4 | 6.7 | 2.4 | 2.6 | 4.9 | 2.5 | 12.2 | 11.9 | 11.9 | 19.0 | 1.5 | ||
ROM-36522 | 11 | m | 6.8 | 6.6 | 2.2 | 2.7 | 4.3 | 2.3 | 12.3 | 12.9 | 11.3 | 18.7 | 1.2 | ||
ROM-36528 | 11 | m | 6.1 | 5.8 | 2.2 | 2.4 | 4.2 | 2.5 | 12.5 | 13.0 | 11.7 | 18.4 | 1.7 | ||
ROM-36526 | 11 | m | 6.1 | 6.3 | 2.3 | 2.5 | 4.3 | 2.7 | 12.7 | 12.4 | 12.7 | 18.2 | 2.2 | ||
8 | m | 7.8 | 7.1 | 3.2 | 2.9 | 4.1 | 3.4 | 4.6 | 12.0 | 11.2 | 19.6 | 1.8 | |||
8 | m | 7.5 | 8.0 | 3.5 | 2.8 | 5.1 | 3.9 | 14.8 | 14.9 | 12.7 | 22.8 | 1.9 | |||
8 | m | 7.8 | 8.2 | 3.1 | 3.8 | 5.2 | 4.1 | 16.3 | 16.1 | 12.6 | 21.9 | 2.1 | |||
8 | m | 7.0 | 7.3 | 3.4 | 3.3 | 5.0 | 3.5 | 19.6 | 13.4 | 12.1 | 20.0 | 1.7 | |||
8 | m | 8.9 | 8.3 | 3.4 | 3.0 | 6.0 | 3.6 | 16.0 | 15.1 | 12.3 | 21.9 | 2.1 | |||
8 | m | 7.8 | 7.7 | 3.2 | 3.0 | 5.5 | 3.5 | 15.0 | 14.7 | 12.6 | 20.7 | 1.8 | |||
ABV-00455 | 6 | f | paratype | 8.3 | 7.7 | 2.9 | 3.3 | 4.9 | 3.1 | 16.1 | 16.2 | 14.2 | 22.6 | 1.7 | |
CYS-10-10 | 11 | f | 7.9 | 8.6 | 3.0 | 3.4 | 5.2 | 3.2 | 15.3 | 15.5 | 13.9 | 21.8 | 2.4 | ||
ROM-36530 | 11 | f | 7.7 | 8.7 | 3.3 | 4.7 | 5.6 | 3.6 | 14.9 | 15.8 | 15.1 | 23.3 | 1.5 | ||
8 | f | 9.8 | 9.2 | 4.1 | 3.9 | 6.3 | 4.6 | 19.8 | 17.8 | 15.7 | 27.2 | 2.3 | |||
8 | f | 10.1 | 9.7 | 4.2 | 4.1 | 7.0 | 4.9 | 17.7 | 15.5 | 15.3 | 25.0 | 2.0 | |||
8 | f | 10.1 | 9.1 | 4.5 | 4.1 | 5.8 | 5.2 | 19.0 | 17.2 | 15.1 | 26.0 | 2.1 |
Tadpoles were allocated to
Standard tadpoles measurements (mean ± SD,
The following description is based on five tadpoles at stage 25 (
External morphology of the tadpole of
In life tadpoles have dorsal side of body and upper flanks uniform brownish red or brownish orange (Fig.
Coloration of
Refer to the Acoustic differentiation section, Table
The new species is reconstructed as a member of the
Comparison of the head coloration in life of three
All specimens were collected at night after heavy rains along montane cascade rocky streams, along small waterfalls, or intermittent rocky brooks; or found during the day time under tree-logs and within leaf litter in the limited fragments of primary montane wet polydominant evergreen tropical forests, with a high abundance of large rocks and fallen trees covered with a thick layer of mosses. This including high montane forests that are composed of the specific floral community known as “elfin” forests, with miniature trees (up to 10 m tall). These areas always have high precipitation and have much milder climate than other tropical forests in southern Vietnam: active breeding of the new species was recorded in February with temperatures of ca. 11–12°C.
On Bidoup Mt. summit (Lam Dong Prov.),
On Bidoup Mt. summit (1890–2035 m a.s.l.; Lam Dong Prov.)
Natural habitat of
Reproductively active males were found while calling along streams, usually sitting on leaves of ferns or on the stone banks, rarely on rocks or large stones (see Fig.
The ovaries of females contained well-developed unpigmented eggs with a diameter of approximately 2.2–2.8 mm (
The full extent of the distribution of
Though available information on tadpole morphology of
DNA-barcoding using short sequences for 16S rRNA (Table
Despite overall similarity, advertisement calls of each
The call temporal parameters for
Finally, the new species is markedly distinct from all other congeners for which comparable sequences are available, including it closest relatives
The data presented here provide the most extensive molecular sampling for the elucidation of phylogentic relationships within the genus
A hidden diversity of
The frequency of maximum amplitude coincides with the fundamental frequency for all
The Langbian Plateau is known for its high herpetofaunal diversity and endemism, a significant portion of which has been discovered only recently (e.g.,
Habitat loss is the greatest threat to amphibians in southeast Asia, and the amphibians of the region appear to be particularly vulnerable to habitat alterations (
Due to the simultaneous review period of the present paper, and the now recently published
NA Poyarkov envisioned the original idea of the manuscript, collected material and data in the field and in the lab, executed this study and wrote the manuscript; TV Duong performed morphometric, molecular and phylogenetic analyses; NL Orlov collected material in the field; SS Gogoleva collected data in the field and performed acoustic analyses and wrote the relevant parts of the manuscript; AB Vassilieva collected material and data in the field; LT Nguyen collected material in the field and assisted with morphological analysis; VDH Nguyen, J Che and
Fieldwork was funded by the Joint Russian-Vietnamese Tropical and Technological Center (JRVTTC) and was conducted under permission of the Bureau of Forestry, Ministry of Agriculture and Rural Development of Vietnam (permits Nos. 170/ TCLN–BTTN of 07/02/2013; 400/TCLN-BTTN of 26/03/2014; 831/TCLN–BTTN of 05/07/2013) and of local administration (Lam Dong Prov.: No. 5832/UBND-LN of 22/10/12; Khanh Hoa Prov.: No. 522/SngV-TTDN&HTQT of 13/07/2013; fieldwork was conducted in accordance to the Agreement No. 37/HD on the scientific cooperation between Cat Tien N.P. and the JRVTTC and the Agreement № 137/HD NCKH of 23.06.2010 on the scientific cooperation between Bu Gia Map N.P. and the JRVTTC). The authors are grateful to Andrei N. Kuznetsov, Leonid P. Korzoun and Vitaly L. Trounov for support and organization of fieldwork. We sincerely thank our Vietnamese colleagues Nguyen Dang Hoi, Hoang Minh Duc, Nguyen Ngoc Hung, Nguyen Thien Tao, Pham Thi Ha Giang, Tran Tien and Le Xuan Son for help and continued support. Many thanks to Eduard A. Galoyan, Igor V. Palko, Evgeniy S. Popov, Olga V. Morozova, Alina V. Alexandrova, Valentina D. Kretova and Evgeniya N. Solovyeva for their help during the fieldwork, assistance in the laboratory and continued support of this project. We thank Ilya A. Volodin for help with acoustic analyses. For permission to study specimens under their care, we thank Valentina F. Orlova (
Examined material, museum IDs given in bold.
Factor coordinates of the morphometric characters used in
Character | factor 1 | factor 2 | factor 3 |
---|---|---|---|
|
-0.969013 | -0.052119 | -0.061729 |
|
-0.972214 | -0.094198 | -0.099264 |
|
-0.963376 | -0.112316 | -0.089300 |
|
-0.872338 | -0.060587 | 0.293135 |
|
-0.853903 | -0.302433 | 0.054377 |
|
-0.869868 | -0.053510 | -0.256344 |
|
-0.817708 | 0.454058 | -0.126722 |
|
-0.774425 | -0.018511 | -0.529049 |
|
-0.302259 | 0.915825 | 0.017344 |
|
-0.866068 | -0.171158 | 0.204049 |
|
-0.818811 | 0.248915 | 0.129748 |
|
-0.848819 | 0.181381 | 0.241783 |
|
-0.937173 | -0.162120 | -0.073309 |
|
-0.957549 | 0.054374 | -0.001476 |
|
-0.931528 | -0.047484 | 0.090017 |
|
-0.939026 | -0.049085 | 0.087852 |
|
-0.963650 | 0.030269 | -0.041650 |
|
-0.902097 | 0.102388 | -0.091818 |
|
-0.950468 | -0.104557 | 0.046962 |
|
-0.921529 | -0.091021 | 0.019328 |
|
-0.918076 | 0.080433 | 0.100931 |
|
-0.971401 | 0.003596 | 0.002474 |
|
-0.761261 | -0.079858 | 0.068715 |
Measurements of advertisement call temporal parameters and one-way ANOVA/Kruskal-Wallis results for comparison (*p < 0.001) between April and February sets of call recordings for
Parameters | Tukey/ |
ANOVA/ |
||
---|---|---|---|---|
Temperature of recording | 11.3–11.4°C | 17.0–17.5°C | – | – |
Number of males | 2 | 1 | – | – |
Number of recordings | 3 | 2 | – | – |
Number of series | 93 | 47 | – | – |
Number of calls | 1301 | 496 | – | – |
Call repetition rate per recording (calls/s) | 1.25 ± 0.35 |
1.07 ± 0.06 |
p = 0.71 | F1.3 = 0.2 |
Number of calls per series | 14 ± 0.45 |
10.53 ± 0.72 |
p < 0.001 | F1.138 = 18.2* |
Series duration (s) | 3.37 ± 0.1 |
3.52 ± 0.27 |
p = 0.52 | F1.138 = 0.4 |
Call repetition rate per series (calls/s) | 4.22 ± 0.08 |
3.18 ± 0.1 |
p < 0.001 | F1.138 = 63.9* |
Call duration (ms) | 70 ± 0.27 |
79 ± 0.32 |
p < 0.001 |
H1.1798 = 380.2* |
Inter-calls interval (ms) | 184 ± 1.72 |
271.17 ± 4.92 |
p < 0.001 |
H1.1657 = 349.2* |
Inter-series interval (s) | 6.72 ± 0.54 |
6.09 ± 0.56 |
p = 0.83 |
H1.136 = 0.05 |