# Phylogeographic pattern and taxonomic revision of the Kaloula baleata species complex (Amphibia, Anura, Microhylidae) with description of two new species from Indochina

Vladislav A Gorin, Nikolai L Orlov, Andrey M Bragin, Parinya Pawangkhanant, Konstantin D Milto, Dac Xuan Le, Tan Van Nguyen, Christophe Dufresnes, Chatmongkon Suwannapoom, Nikolay A Poyarkov

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Vladislav A Gorin, Nikolai L Orlov, Andrey M Bragin, Parinya Pawangkhanant, Konstantin D Milto, et al.. Phylogeographic pattern and taxonomic revision of the Kaloula baleata species complex (Amphibia, Anura, Microhylidae) with description of two new species from Indochina. Herpetozoa, 2024, 37, pp.391-420. 10.3897/herpetozoa.37.e137394. hal-04886307

HAL Id: hal-04886307

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# Phylogeographic pattern and taxonomic revision of the *Kaloula baleata* species complex (Amphibia, Anura, Microhylidae) with description of two new species from Indochina

Vladislav A. Gorin¹, Nikolai L. Orlov², Andrey M. Bragin³, Parinya Pawangkhanant⁴,⁵, Konstantin D. Milto², Dac Xuan Le³, Tan Van Nguyen⁶,⁷, Christophe Dufresnes⁸, Chatmongkon Suwannapoom⁴, Nikolay A. Poyarkov¹,³

1 Department of Vertebrate Zoology, Lomonosov Moscow State University, Leninskiye Gory, Moscow 119234, Russia

2 Division of Herpetology & Ornthology, Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034 Universitetskaya nab., 1, 33701, Russia

3 Joint Vietnam - Russia Tropical Science & Technology Research Center, 63 Nguyen Van Huyen Road, Nghia Do, Cau Giay, Hanoi 122000, Vietnam

4 Division of Fishery, School of Agriculture & Natural Resources, University of Phayao, Phayao 56000, Thailand

5 Rabbit in the Moon Foundation, Suanphuang, Ratchaburi 70180, Thailand

6 Institute for Research & Training in Medicine, Biology & Pharmacy, Duy Tan University, Da Nang 550000, Vietnam

7 College of Medicine & Pharmacy, Duy Tan University, 120 Hoang Minh Thao, Lien Chieu, Da Nang 550000, Vietnam

8 Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS, Sorbonne Université, EPHE-PSL, Université des Antilles, Paris, France

https://zoobank.org/E97A128A-13B5-4781-8FEC-ADC5CB37AB45

Corresponding authors: Nikolay A. Poyarkov (n.poyarkov@gmail.com); Chatmongkon Suwannapoom (chatmongkonup@gmail.com)

Academic editor: Peter Mikulíček ♦ Received 18 September 2024 ♦ Accepted 22 November 2024 ♦ Published 11 December 2024

## Abstract

Despite increased attention by molecular taxonomists, the herpetofauna of Southeast Asia still hides many undescribed species among far-ranging taxa. In this study, we re-examine the mitochondrial diversity of painted frogs of the microhylid genus *Kaloula*, based on ~2,455 bp of published and new 12S and 16S rRNA sequences, and describe two new species from the tropical forests of southern Vietnam and central Laos based on integrative evidence. These species, which belong to the *K. baleata* complex, feature species-level mitochondrial divergence (> 4.4% at 16S rRNA) and are both morphologically well-differentiated from each other and from the recently described *K. indochinensis*, to which they were previously confounded. Comparative examinations also indicate distinct male advertisement calls and unique coloration features. Based on genetic barcoding, we preliminarily revise the species distribution ranges in the *K. baleata* complex, which support a general pattern of biogeographic partitioning that has been widely retrieved among the Indochinese amphibians studied so far. Molecular diversity within *K. baleata* further suggests genetic structure across Sundaland, namely three shallow mitochondrial haplogroups worthy of fine-scale phylogeographic and taxonomic investigations. Furthermore, our study highlights the propensity of amphibian species “hidden in plain sight,” even among recently studied taxa, thus calling for caution when specifying type specimens—the type series of *K. indochinensis*, described in 2013, includes specimens of the one of the new species. Our study emphasizes the continued need for thorough herpetological surveys even in supposedly well-known parts of Indochina and sets the ground for future research in *Kaloula* painted frogs, notably to test evolutionary and taxonomic hypotheses with genomic loci.

## Key Words

16S rRNA, bioacoustics, DNA barcoding, Laos, morphology, Southeast Asia, systematics, taxonomy, Vietnam
# Introduction

Many widely distributed amphibian genera in Asia have undergone a considerable increase in their numbers of recognized species since the turn of the 21st century. For example, among microhylids of the genus *Microhyla* Tschudi, 1838, the number of recognized species jumped from 22 to 51 within two decades (Poyarkov et al. 2020a; Zhang et al. 2022; Trofimets et al. 2024). The number of species within the genus *Micryletta* Dubois, 1987, rose from three in the 20th century to 13 species nowadays (Suwannapoom et al. 2020; Poyarkov et al. 2021a; Sankar et al. 2022). The genus *Kalophrynus* Tschudi, 1838, included from 13 to currently 27 recognized species (Fukuyama et al. 2021). Two new microhylid genera were discovered in Southeast Asia in the past several years, namely, *Siamophryne* Suwannapoom, Su-montha, Tunprasert, Ruangsuwan, Pawangkhanant, Korost & Poyarkov, 2018 (one single known species) and *Viet-namophryne* Poyarkov, Suwannapoom, Pawangkhanant, Aksornneam, Duong, Korost & Che, 2018 (six species described so far; see Suwannapoom et al. 2018; Poyarkov et al. 2018a, 2021b; Gorin et al. 2021; Frost 2024).

Quite a few genera of Asian microhylids, however, have so far evaded the focus of taxonomic studies, and their species numbers thus remained relatively stable. One notable example is the genus *Kaloula* Gray, 1831, also known as Asian painted frogs (Frost 2024). *Kaloula* is a widely distributed genus of microhylids, with 18 species recognized up to date. Painted frogs inhabit East Asia, from Korea and northeastern China to Southeast Asia, including Indochina to the Sunda Islands and the Philippines to the south, and Bangladesh and northeastern India to the west (Othman et al. 2022; Frost 2024). The genus *Kaloula* is characterized by the following morphological attributes: pupil round; tongue oval, entire, and free behind; two transverse ridges across the palate in front of the pharynx; a strong bony ridge behind each choanae; the first finger shorter than the second and the outer toe shorter than the third; medium body size (SVL 35–60 mm); smooth or slightly rough dorsum with irregular dark markings; tips of the fingers dilated into disks and truncated; fingers free of the web; nearly full webbing on toes in males and reduced webbing in females (e.g., Gray 1831; Sengupta et al. 2009; Gorin et al. 2021). Recent contributions to the diversity of this genus include the review by Blackburn et al. (2013), which focused on *Kaloula* species inhabiting Southeast Asian islands, mainly the Philippine Archipelago. In this study, the authors conducted phylogenetic analysis on 140 *Kaloula* individuals from all over Southeast Asia and indicated the presence of at least six undescribed lineages of putative species status. Following the study of Blackburn et al. (2013), two new species of the *K. baleata* complex were described from Indochina, namely *K. indochinensis* Chan, Blackburn, Murphy, Stuart, Emmett, Ho & Brown, 2013 (Chan et al. 2013), and from Peninsular Malaysia, namely *K. latidisca* Chan, Grismer & Brown, 2014 (Chan et al. 2014). In both cases, authors provided only limited

morphological data to substantiate species divergence, did not provide any bioacoustic data, and based their phylogenetic estimations entirely on the previously published mtDNA data of Blackburn et al. (2013).

In parallel, a new species, *K. nonggangensis* Mo, Zhang, Zhou, Chen, Tang, Meng & Chen, 2013, was described from the Guangxi Province of China (Mo et al. 2013) on the basis of morphological and genetic data. More recently, the subspecies *K. baleata ghoshi* Cherchi, 1954, endemic to the Andaman Islands, was elevated to full species status based on morphological differentiation, accompanied by new data on natural history, bioacoustics, and distribution (Chandramouli and Prasad 2018). Lastly, the phylogeography of the genus *Kaloula* in East Asia was re-summarized by Ohtman et al. (2022), based mostly on previously published data, and without emphasizing undescribed diversity.

In this work, we present an updated mitochondrial DNA (hereafter mtDNA)-based genealogy for the genus *Kaloula*, combining sequences available in GenBank with new sequences obtained from an extensive sampling in Indochina, as well as integrative taxonomic revisions in the *K. baleata* species complex, specifically for two populations from central Laos and southern Vietnam previously assigned to *K. indochinensis* (Chan et al. 2013). These populations are deeply divergent in the mitochondrial phylogeny and feature morphological and bioacoustic differentiation from related taxa, hence substantiating their description as new species.

# Materials and methods

## Sample collection

Fieldwork in Vietnam was conducted by NAP, AMB, and VAG in 2009–2023; fieldwork in Laos was conducted by NLO and KDM in 2009. We obtained geographic coordinates and elevation data using a Garmin GPSMAP 60CSx (USA) and recorded them in the WGS84 datum. The specimens collected were initially fixed in 4% formalin for 24 h and later transferred to 75% ethanol for storage; muscle or liver tissues were taken prior to the fixation in formalin and preserved in 95% ethanol for genetic analysis. Specimens were subsequently deposited in the herpetological collections of the Zoological Museum of Lomonosov Moscow State University (ZMMU, Moscow, Russia) and of the Zoological Institute, Russian Academy of Sciences (ZISP, St. Petersburg, Russia).

## Laboratory methods

For the molecular phylogenetic analyses, total genomic DNA was isolated using the standard phenol-chloroform-proteinase K extraction procedures with consequent isopropanol precipitation for a final concentration of about 1 mg/ml (protocols followed the works of Hillis et
al. 1996; Sambrook and Russell 2001). We visualized the isolated total genomic DNA using agarose electrophoresis in the presence of ethidium bromide. We measured the concentration of total DNA in 1 µL using NanoDrop 2000 (Thermo Scientific) and consequently adjusted it to ca. 100 ng DNA/µL.

We amplified mtDNA fragments covering the 16S rRNA gene and the adjacent tRNA genes to obtain a 1,003 bp-long continuous fragment. The 16S rRNA gene is widely used for biodiversity surveys in amphibians (Vences et al. 2005; Vieites et al. 2009) and has been analyzed in the most recent phylogenetic studies on Microhylinae (Matsui et al. 2011; Garg et al. 2019; Gorin et al. 2020, 2021). We performed DNA amplification in 20 µL reactions using ca. 50 ng genomic DNA, 10 nmol of each primer, 15 nmol of each dNTP, 50 nmol additional MgCl₂, Taq PCR buffer (10 mM Tris-HCl, pH 8.3, 50 mM KCl, 1.1 mM MgCl₂, and 0.01% gelatine), and 1 unit of Taq DNA polymerase. Primers used in PCR and sequencing include 16sL-2188 (CTGACCGTGCAAAGGTAGCGTAATCACT) and 16H-1 (CTCCGGTCTGAACTCAGATCACGTAGG) (Hedges et al. 1994; Matsui et al. 2005). The PCR conditions involved an initial denaturation step of 5 min at 94 °C, followed by 43 cycles of denaturation for 1 min at 94 °C, primer annealing for 1 min using the TouchDown program, reducing 1 °C every cycle, extension for 1 min at 72 °C, and a final extension step for 5 min at 72 °C (Gorin et al. 2020, 2021).

PCR products were loaded onto 1.0% agarose gels in the presence of ethidium bromide and visualized in agarose electrophoresis. When distinct bands were obtained, we purified PCR products using 2 µL of a 1:4 dilution of ExoSapIt (Amersham) per 5 µL of PCR product prior to cycle sequencing. A 10 µL sequencing reaction included 2 µL of template, 2.5 µL of sequencing buffer, 0.8 µL of 10 pmol primer, 0.4 µL of BigDye Terminator version 3.1 Sequencing Standard (Applied Biosystems), and 4.2 µL of water. The cycle sequencing used 35 cycles of 10 sec at 96 °C, 10 s at 50 °C, and 4 min at 60 °C. We purified the cycle sequencing products by ethanol precipitation. We carried out sequence data collection and visualization on an ABI 3730xl Automated Sequencer (Applied Biosystems). We deposited the obtained sequences in GenBank under the accession numbers PQ268497–PQ268520.

## Phylogenetic analyses

To reconstruct the matrilineal genealogy, we used newly obtained 16S rRNA sequences of *Kaloula* spp. from Laos and Vietnam and the sequences of 12S rRNA and 16S rRNA mtDNA fragments of the *K. baleata* species complex members from Thailand, Malaysia, and Indonesia, as well as other *Kaloula* species obtained from GenBank. Table 1 summarizes the information on GenBank Accession Numbers, museum vouchers, and the locality of origin for the sequences used in this study. We also added sequences of *Uperodon taprobanicus* (Parker, 1934) as a sister group;

a sequence of *Kalophrynus interlineatus* (Blyth, 1855) was used to root the tree; the final alignment included 2,455 bp. In total, we obtained data for 16S rRNA for 233 specimens, which included 15 out of 18 currently recognized *Kaloula* species, including the topotype specimens of *K. baleata* (Müller, 1836) (type locality: Java, Indonesia) (see Table 1); geographic distribution of the sampled populations of the *K. baleata* species complex is shown in Fig. 1.

We initially aligned nucleotide sequences using ClustalX 1.81 (Thompson et al. 1994) with default parameters and then optimized them manually in BioEdit 7.0.5.2 (Hall 1999) and MEGA 11.0 (Tamura et al. 2013). We utilized ModelFinder (Kalyaanamoorthy et al. 2017) to determine the most suitable evolutionary models for our data set analysis. According to the Akaike Information Criterion (AIC), the best-fitting models of DNA evolution for 12S rRNA, tRNA-Val, and 16S rRNA genes of mtDNA are GTR+I+G, K2P+G, and GTR+I+G, respectively. We determined mean uncorrected genetic distances (p-distances) between sequences with MEGA 11.0.

We inferred the matrilineal genealogy using Bayesian Inference (BI) and Maximum Likelihood (ML) approaches. We conducted BI using MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). Metropolis-coupled Markov chain Monte Carlo (MCMCMC) analyses were run with one cold chain and three heated chains for one million generations and sampled every 1,000 generations. We performed two independent MCMCMC runs, and the initial 100 trees were discarded as burn-in. We assessed confidence in tree topology based on the frequency of nodal resolution (posterior probability; BI PP) (Huelsenbeck and Ronquist 2001). We used IQ-TREE (Nguyen et al. 2015) to reconstruct ML-trees. A total of 10,000 ultrafast bootstrap replications for ML analysis (UFB) (Minh et al. 2013) assessed the confidence in tree topology for ML analysis. In both datasets, we regarded tree nodes with BI PP and UFB values over 0.95 to be sufficiently resolved a priori. We considered BI PP and UFB values between 0.95 and 0.90 as tendencies. Lower values were considered to indicate unresolved nodes (Huelsenbeck and Hillis 1993; Minh et al. 2013).

## Morphological analysis

Measurements were taken using a digital caliper under a light dissecting microscope to the nearest 0.01 mm, subsequently rounded to 0.1 mm. The adult morphometrics and character terminology follow Poyarkov et al. (2018b): (1) snout-vent length (SVL; measured from the tip of the snout to cloaca); (2) head length (HL; measured from the tip of snout to hind border of jaw angle); (3) snout length (SL; measured from the anterior corner of eye to the tip of snout); (4) eye length (EL; measured as the distance between anterior and posterior corners of the eye); (5) nostril-eye length (N-EL; measured as the distance between the anterior corner of the eye and the nostril center); (6) head width (HW; measured as the maximum width of head on the level of mouth angles in ventral view); (7) internarial
<figure><img src="image_2.png" /><figcaption>**Figure 1.** Geographic origins of the specimens of the *Kaloula baleata* species complex included in phylogenetic analyses. Dots in the symbols denote type localities. Symbol colors correspond to those used in Fig. 2. Numbers near symbols correspond to the locality numbers given in Table 1. The white rectangle represents the area of inset map detailing the distribution of the *K. baleata* species complex in Indochina in Fig. 5.</figcaption></figure>

distance (IND; measured as the distance between the central points of nostrils); (8) interorbital distance (IOD; measured as the shortest distance between the medial edges of eyeballs in dorsal view); (9) upper eyelid width (UEW; measured as the maximum distance between the medial edge of eyeball and the lateral edge of upper eyelid); (10) tympanum length, measured as the horizontal tympanum diameter (TMP); (11) forelimb length (FLL; measured as the length of straightened forelimb to the tip of third finger); (12) lower arm and hand length (LAL; measured as the distance between elbow and the tip of third finger); (13) hand length (HAL; measured as the distance between the proximal end of outer palmar (metacarpal) tubercle and the tip of third finger); (14) first finger length (1FL, measured as the distance between the tip and the distal end of inner palmar tubercle); (15) inner palmar tubercle length (IPTL; measured as the maximum distance between proximal and

distal ends of inner palmar tubercle); (16) outer palmar tubercle length (OPTL; measured as the maximum diameter of outer palmar tubercle); (17) third finger disk diameter (3FDD); (18) hindlimb length (HLL; measured as the length of straightened hindlimb from groin to the tip of fourth toe); (19) tibia length (TL; measured as the distance between the knee and tibiotarsal articulation); (20) foot length (FL; measured as the distance between the distal end of tibia and the tip of fourth toe); (21) first toe length (1TOEL), measured as the distance between the distal end of inner metatarsal tubercle and the tip of first toe; (22) fourth toe disk diameter (4TDD); (23) outer metatarsal tubercle length (OMTL). Webbing and subarticular tubercle formulas follow those of Savage (1975). We took all measurements on the right side of the examined specimen. We determined the sex of specimens by examining the presence of eggs in the abdomen of a dissected specimen.
<table><caption>**Table 1.** Museum voucher information, geographic localities, and GenBank accession numbers of specimens and sequences used in this study. Exact locality information is unknown for specimens obtained via the pet trade or those published in some earlier works. Locality numbers given in brackets in bold after the locality name correspond to the numbers shown in Fig. 1.</caption><thead><tr><th rowspan="2">No.</th><th rowspan="2">Species<br>Ingroup</th><th rowspan="2">Locality</th><th rowspan="2">Museum / Sample ID</th><th colspan="2">Accession numbers</th><th rowspan="2">Reference</th></tr><tr><th>12S rRNA</th><th>16S rRNA</th></tr></thead><tbody><tr><td colspan="7"><strong>Kaloula baleata species complex</strong></td></tr><tr><td>1</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Dong Nai Prov., Cat Tien NP (1)</td><td>ZMMU-A-4642</td><td></td><td>PQ268510</td><td>this work</td></tr><tr><td>2</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Dong Nai Prov., Cat Tien NP (1)</td><td>ZMMU-A-4739</td><td></td><td>PQ268511</td><td>this work</td></tr><tr><td>3</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Dong Nai Prov., Cat Tien NP (1)</td><td>ZISP 15285</td><td></td><td>PQ268512</td><td>this work</td></tr><tr><td>4</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Dong Nai Prov., Cat Tien NP (1)</td><td>ZMMU-A-8134</td><td></td><td>PQ268513</td><td>this work</td></tr><tr><td>5</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Dong Nai Prov., Cat Tien NP (1)</td><td>ZMMU-A-8135</td><td></td><td>PQ268514</td><td>this work</td></tr><tr><td>6</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Lam Dong Prov., Loc Bac (2)</td><td>ZMMU NAP-02826</td><td></td><td>PQ268515</td><td>this work</td></tr><tr><td>7</td><td>Kaloula discordia sp. nov.</td><td>Vietnam, Lam Dong Prov., Loc Bac (2)</td><td>ZMMU NAP-03384</td><td></td><td>PQ268516</td><td>this work</td></tr><tr><td>8</td><td>Kaloula laosensis sp. nov.</td><td>Laos, Khammouane Prov., Nakai-Nam Theun NP (3)</td><td>ZISP 15284</td><td></td><td>PQ268517</td><td>this work</td></tr><tr><td>9</td><td>Kaloula laosensis sp. nov.</td><td>Laos, Khammouane Prov., Nakai-Nam Theun NP (3)</td><td>ZMMU-A-8144</td><td></td><td>PQ268518</td><td>this work</td></tr><tr><td>10</td><td>Kaloula indochinensis</td><td>Vietnam, Dak Lak Prov., Yok Don NP (4)</td><td>ZMMU-A-8147</td><td></td><td>PQ268497</td><td>this work</td></tr><tr><td>11</td><td>Kaloula indochinensis</td><td>Vietnam, Dak Lak Prov., Yok Don NP (4)</td><td>ZMMU-A-8148</td><td></td><td>PQ268498</td><td>this work</td></tr><tr><td>12</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ROM32925</td><td></td><td>KC180032</td><td>de Sa et al. 2012</td></tr><tr><td>13</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ZISP-Gia Lai 1</td><td></td><td>PQ268499</td><td>this work</td></tr><tr><td>14</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ZISP-Gia Lai 2</td><td></td><td>PQ268500</td><td>this work</td></tr><tr><td>15</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ROM32925</td><td>KC822572</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>16</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ROM32932</td><td>KC822573</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>17</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Krong Pa (5)</td><td>ROM32943</td><td>KC822574</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>18</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Kon Ka Kinh NP (6)</td><td>ZMMU-A-6315</td><td></td><td>PQ268501</td><td>this work</td></tr><tr><td>19</td><td>Kaloula indochinensis</td><td>Vietnam, Gia Lai Prov., Kon Ka Kinh NP (6)</td><td>ZMMU-A-6316</td><td></td><td>PQ268502</td><td>this work</td></tr><tr><td>55</td><td>Kaloula latidisca</td><td>Malaysia, Kedah, Gubir (7)</td><td>LSUHC5074</td><td>KC822576</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>56</td><td>Kaloula latidisca</td><td>Thailand, Satun Prov., Baan Suan Tondin (8)</td><td>ZMMU-A-8145</td><td></td><td>PQ268503</td><td>this work</td></tr><tr><td>57</td><td>Kaloula latidisca</td><td>Thailand, Surathani Prov., Khao Sok (9)</td><td>ZMMU-A-8146</td><td></td><td>PQ268504</td><td>this work</td></tr><tr><td>20</td><td>Kaloula baleata clade 1</td><td>Indonesia, Java isl., Jawa Barat (10)</td><td>TNHC67086</td><td>KC822569</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>21</td><td>Kaloula baleata clade 1</td><td>Indonesia, Bali Prov., Bali isl. (11)</td><td>JAM3232</td><td>KC822570</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>22</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sumatra, Way Jepara (12)</td><td>ZMMU A-04412</td><td></td><td>PQ268507</td><td>this work</td></tr><tr><td>23</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sumba (13)</td><td>KUHE32313</td><td>AB634629</td><td>AB634687</td><td>Matsui et al. 2011</td></tr><tr><td>24</td><td>Kaloula baleata clade 1</td><td>Philippines, Palawan isl. (14)</td><td>ACD1303</td><td>KC822582</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>25</td><td>Kaloula baleata clade 1</td><td>Philippines, Palawan isl. (14)</td><td>ACD1307</td><td>KC822583</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>26</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sulawesi (-)</td><td>JAM3573</td><td></td><td>KY132185</td><td>Alexander et al. 2017</td></tr><tr><td>27</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sulawesi Tengah Prov., Pulau Batudaka (15)</td><td>JAM-3853</td><td></td><td>KC822568</td><td>Blackburn et al. 2013</td></tr><tr><td>28</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sulawesi Tengah Prov., Donggala (16)</td><td>LSUMZ-83998</td><td></td><td>KC822566</td><td>Blackburn et al. 2013</td></tr><tr><td>29</td><td>Kaloula baleata clade 1</td><td>Indonesia, Sulawesi Utara Prov., Bogani Nani<br>Wartabone NP (17)</td><td>LSUMZ-83999</td><td></td><td>KC822567</td><td>Blackburn et al. 2013</td></tr><tr><td>30</td><td>Kaloula baleata clade 2</td><td>Malaysia (-)</td><td>FRIM1066</td><td>KC822579</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>31</td><td>Kaloula baleata clade 2</td><td>Malaysia (-)</td><td>DWNP975</td><td>KC822580</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>32</td><td>Kaloula baleata clade 2</td><td>Malaysia, Johor, Pulau Aceh (18)</td><td>LSUHC5712</td><td>KC822577</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>33</td><td>Kaloula baleata clade 2</td><td>Malaysia, Sabah, Sepilok (19)</td><td>LSUHC6156</td><td>KC822578</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>35</td><td>Kaloula baleata clade 2</td><td>Malaysia, Sabah, Kinabalu (20)</td><td>ZMHA10028</td><td></td><td>GU154880</td><td>Das and Haas 2010</td></tr><tr><td>34</td><td>Kaloula baleata clade 2</td><td>Malaysia, Sarawak, Kuching (21)</td><td>ID8317</td><td>KC822581</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>36</td><td>Kaloula baleata clade 2</td><td>Malaysia, Pahang, Taman Negara (22)</td><td>ZMMU-A-6152</td><td></td><td>PQ268508</td><td>this work</td></tr><tr><td>37</td><td>Kaloula baleata clade 2</td><td>Malaysia, Pahang, Taman Negara (22)</td><td>ZMMU-A-6153</td><td></td><td>PQ268509</td><td>this work</td></tr><tr><td>38</td><td>Kaloula baleata clade 2</td><td>Indonesia, Java, Bogor (23)</td><td>released (Java)</td><td></td><td>PQ268505</td><td>this work</td></tr><tr><td>39</td><td>Kaloula baleata clade 2</td><td>Indonesia, Java, Bogor (23)</td><td>released (Java)</td><td></td><td>PQ268506</td><td>this work</td></tr><tr><td>40</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587865</td><td></td><td>MT608909</td><td>Mulcahy et al. unpublished</td></tr><tr><td>41</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587869</td><td></td><td>MT608910</td><td>Mulcahy et al. unpublished</td></tr><tr><td>42</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587870</td><td></td><td>MT608911</td><td>Mulcahy et al. unpublished</td></tr><tr><td>43</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587871</td><td></td><td>MT608912</td><td>Mulcahy et al. unpublished</td></tr><tr><td>44</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587602</td><td></td><td>MT608913</td><td>Mulcahy et al. unpublished</td></tr><tr><td>45</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587603</td><td></td><td>MT608914</td><td>Mulcahy et al. unpublished</td></tr><tr><td>46</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587863</td><td></td><td>MT608915</td><td>Mulcahy et al. unpublished</td></tr><tr><td>47</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587864</td><td></td><td>MT608916</td><td>Mulcahy et al. unpublished</td></tr><tr><td>48</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587866</td><td></td><td>MT608917</td><td>Mulcahy et al. unpublished</td></tr></tbody></table>
<table><thead><tr><th rowspan="2">No.</th><th>Species</th><th rowspan="2">Locality</th><th rowspan="2">Museum / Sample ID</th><th colspan="2">Accession numbers</th><th rowspan="2">Reference</th></tr><tr><th>Ingroup</th><th>12S rRNA</th><th>16S rRNA</th></tr></thead><tbody><tr><td>49</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587867</td><td>MT608918</td><td></td><td>Mulcahy et al. unpublished</td></tr><tr><td>50</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587868</td><td>MT608919</td><td></td><td>Mulcahy et al. unpublished</td></tr><tr><td>51</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587604</td><td>MT608920</td><td></td><td>Mulcahy et al. unpublished</td></tr><tr><td>52</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587861</td><td>MT608921</td><td></td><td>Mulcahy et al. unpublished</td></tr><tr><td>53</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-587862</td><td>MT608922</td><td></td><td>Mulcahy et al. unpublished</td></tr><tr><td>54</td><td>Kaloula baleata clade 3</td><td>Myanmar, Tanintharyi Div., Yeybu (24)</td><td>USNM-586944</td><td>MG935845</td><td></td><td>Mulcahy et al. 2018</td></tr><tr><td colspan="7"><strong>Other Kaloula species</strong></td></tr><tr><td>58</td><td>Kaloula borealis</td><td>South Korea, Jeju isl.</td><td>KUHE-33139</td><td>AB634688</td><td></td><td>Matsui et al. 2011</td></tr><tr><td>59</td><td>Kaloula borealis</td><td>South Korea, Jeju isl.</td><td>NIBRAM-000123</td><td>JQ815297</td><td></td><td>Jeong et al. 2013</td></tr><tr><td>60</td><td>Kaloula borealis</td><td>South Korea, Jeju isl.</td><td>NIBRAM-100348</td><td>JQ815298</td><td></td><td>Jeong et al. 2013</td></tr><tr><td>61</td><td>Kaloula borealis</td><td>-</td><td>-</td><td>JQ692869</td><td></td><td>Hwang and Lee, 2012</td></tr><tr><td>62</td><td>Kaloula borealis</td><td>China, Beijing, Xiangshan</td><td>KIZHERP-0173</td><td>JX678908</td><td></td><td>Li et al. 2012</td></tr><tr><td>63</td><td>Kaloula borealis</td><td>China, Beijing, Xiangshan</td><td>KIZHERP-0174</td><td>JX678909</td><td></td><td>Li et al. 2012</td></tr><tr><td>64</td><td>Kaloula conjuncta</td><td>Philippines, Negros isl.</td><td>RMB 2252, PNM / CMNH</td><td>AY326064</td><td></td><td>Darst and Canatella, 2003</td></tr><tr><td>65</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Palola</td><td>ACD 769</td><td>KC822537</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>66</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Bukidnon Prov.</td><td>ACD 996</td><td>KC822536</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>67</td><td>Kaloula conjuncta</td><td>Philippines, Semirara isl., Antique Prov.</td><td>KU301849</td><td>KC822532</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>68</td><td>Kaloula conjuncta</td><td>Philippines, Semirara isl., Antique Prov.</td><td>KU301854</td><td>KC822533</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>69</td><td>Kaloula conjuncta</td><td>Philippines, Polillo isl., Quezon Prov.</td><td>KU303279</td><td>KC822527</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>70</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Laguna Prov.</td><td>KU320031</td><td>KC822538</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>71</td><td>Kaloula conjuncta</td><td>Philippines, Mindoro isl., Mindoro Occidental Prov.</td><td>KU323280</td><td>KC822535</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>72</td><td>Kaloula conjuncta</td><td>Philippines, Mindoro isl., Mindoro Oriental Prov.</td><td>RMB509</td><td>KC822534</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>73</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Quezon Prov.</td><td>TNHC59628</td><td>KC822539</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>74</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62972</td><td>KC822528</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>75</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62973</td><td>KC822526</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>76</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62975</td><td>KC822530</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>77</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62976</td><td>KC822529</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>78</td><td>Kaloula conjuncta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62986</td><td>KC822531</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>79</td><td>Kaloula conjuncta</td><td>Philippines, Camigun Sur isl., Camiguin Sur Prov.</td><td>KU309658</td><td>KC822588</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>80</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59632</td><td>KC822521</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>81</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59635</td><td>KC822520</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>82</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59636</td><td>KC822522</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>83</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59669</td><td>KC822524</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>84</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59670</td><td>KC822523</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>85</td><td>Kaloula conjuncta</td><td>Philippines, Mindanao isl., Davao City Prov.</td><td>TNHC59870</td><td>KC822525</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>86</td><td>Kaloula conjuncta</td><td>Philippines, Negros isl., Negros Oriental Prov.</td><td>KU328639</td><td>KC822518</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>87</td><td>Kaloula conjuncta</td><td>Philippines, Negros isl., Negros Oriental Prov.</td><td>KU328640</td><td>KC822519</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>88</td><td>Kaloula kalingensis</td><td>Philippines, Palaui isl., Cagayan Prov.</td><td>KU328643</td><td>KC822603</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>89</td><td>Kaloula kalingensis</td><td>Philippines, Palaui isl., Cagayan Prov.</td><td>KU328644</td><td>KC822604</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>90</td><td>Kaloula kalingensis</td><td>Philippines, Luzon isl., Kalinga Prov.</td><td>TNHC60118</td><td>KC822605</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>91</td><td>Kaloula kalingensis</td><td>Philippines, Luzon isl., Kalinga Prov.</td><td>RMB2210</td><td>KC822608</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>92</td><td>Kaloula kalingensis</td><td>Philippines, Luzon isl., Kalinga Prov.</td><td>RMB3137</td><td>KC822609</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>93</td><td>Kaloula kalingensis</td><td>Philippines, Luzon isl., Kalinga Prov.</td><td>TNHC59647</td><td>KC822610</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>94</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Camarines del Norte Prov.</td><td>KU313824</td><td>KC822596</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>95</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Camarines Sur Prov.</td><td>KU328629</td><td>KC822595</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>96</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Albay Prov.</td><td>KU328630</td><td>KC822590</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>97</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Quezon Prov.</td><td>KU328634</td><td>KC822593</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>98</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Camarines Sur Prov.</td><td>TNHC62684</td><td>KC822594</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>99</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62685</td><td>KC822589</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>100</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62687</td><td>KC822591</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>101</td><td>Kaloula kokacii</td><td>Philippines, Luzon isl., Sorsogon Prov.</td><td>TNHC62688</td><td>KC822592</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>102</td><td>Kaloula mediolineata</td><td>Thailand, Tak Prov.</td><td>KUHE35178</td><td>AB634631 AB634689</td><td></td><td>Matsui et al. 2011</td></tr><tr><td>103</td><td>Kaloula mediolineata</td><td>Vietnam, Gia Lai Prov.</td><td>ROM32838</td><td>KC822571</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>104</td><td>Kaloula mediolineata</td><td>Thailand, Nakhorn Ratchasima Prov.</td><td>KU328285</td><td>KC822508</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>105</td><td>Kaloula mediolineata</td><td>Thailand, Nakhorn Ratchasima Prov.</td><td>KU328280</td><td>KC822509</td><td></td><td>Blackburn et al. 2013</td></tr></tbody></table>
<table><thead><tr><th rowspan="2">No.</th><th>Species</th><th rowspan="2">Locality</th><th rowspan="2">Museum / Sample ID</th><th colspan="2">Accession numbers</th><th rowspan="2">Reference</th></tr><tr><th>Ingroup</th><th>12S rRNA</th><th>16S rRNA</th></tr></thead><tbody><tr><td>106</td><td>Kaloula mediolineata</td><td>Thailand, Ubon Ratchatani</td><td>FMNH265819</td><td>KC822510</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>107</td><td>Kaloula nonggangensis</td><td>China, Guanxi Prov., Nonggang</td><td>NHMG-1106040</td><td></td><td>KC567231</td><td>Chen et al. 2013</td></tr><tr><td>108</td><td>Kaloula nonggangensis</td><td>China, Guanxi Prov., Nonggang</td><td>NHMG-1108035</td><td></td><td>KC567232</td><td>Chen et al. 2013</td></tr><tr><td>109</td><td>Kaloula nonggangensis</td><td>China, Guanxi Prov., Nonggang</td><td>NHMG-1108036</td><td></td><td>KC567233</td><td>Chen et al. 2013</td></tr><tr><td>110</td><td>Kaloula nonggangensis</td><td>China, Guanxi Prov., Nonggang</td><td>NHMG-CHN-T20120901</td><td></td><td>KC567234</td><td>Chen et al. 2013</td></tr><tr><td>111</td><td>Kaloula picta</td><td>–</td><td>KUHE UN</td><td>AB634686</td><td></td><td>Matsui et al. 2011</td></tr><tr><td>112</td><td>Kaloula picta</td><td>–</td><td>USFS56931</td><td></td><td>KC180019</td><td>de Sa et al. 2012</td></tr><tr><td>113</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Cavite Prov.</td><td>DLSUD021</td><td>KP298039</td><td></td><td>Brown et al. 2015</td></tr><tr><td>114</td><td>Kaloula picta</td><td>Philippines, Palawan isl., Palawan Prov.</td><td>ACD1206</td><td>KC822541</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>115</td><td>Kaloula picta</td><td>Philippines, Leyte isl., Leyte Prov.</td><td>ACD1242</td><td>KC822563</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>116</td><td>Kaloula picta</td><td>Philippines, Palawan isl., Palawan Prov.</td><td>ACD1261</td><td>KC822542</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>117</td><td>Kaloula picta</td><td>Philippines, Palawan isl., Palawan Prov.</td><td>ACD1304</td><td>KC822540</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>118</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Pampanga Prov.</td><td>ACD1390</td><td>KC822546</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>119</td><td>Kaloula picta</td><td>Philippines, Leyte isl., Leyte Prov.</td><td>ACD1501</td><td>KC822562</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>120</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Cavite Prov.</td><td>DLSUD020</td><td>KC822552</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>121</td><td>Kaloula picta</td><td>Philippines, Camiguin Sur Isl., Camiguin Sur Prov.</td><td>KU301873</td><td>KC822565</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>122</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Laguna Prov.</td><td>KU326261</td><td>KC822554</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>123</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Laguna Prov.</td><td>KU326262</td><td>KC822555</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>124</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Quezon Prov.</td><td>KU326265</td><td>KC822548</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>125</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Cagayan Prov.</td><td>RMB4223</td><td>KC822551</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>126</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Cagayan Prov.</td><td>RMB4224</td><td>KC822550</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>127</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Leyte Prov.</td><td>RMB4294</td><td>KC822564</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>128</td><td>Kaloula picta</td><td>Philippines, Leyte isl., Leyte Prov.</td><td>RMB4344</td><td>KC822561</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>129</td><td>Kaloula picta</td><td>Philippines, Bohol isl., Bohol Prov.</td><td>TNHC56430</td><td>KC822559</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>130</td><td>Kaloula picta</td><td>Philippines, Bohol isl., Bohol Prov.</td><td>TNHC56431</td><td>KC822560</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>131</td><td>Kaloula picta</td><td>Philippines, Cebu isl., Cebu Prov.</td><td>TNHC56468</td><td>KC822543</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>132</td><td>Kaloula picta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59653</td><td>KC822558</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>133</td><td>Kaloula picta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59654</td><td>KC822557</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>134</td><td>Kaloula picta</td><td>Philippines, Mindanao isl., Davao del Sur Prov.</td><td>TNHC59871</td><td>KC822556</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>135</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Albay Prov.</td><td>TNHC62470</td><td>KC822549</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>136</td><td>Kaloula picta</td><td>Philippines, Luzon isl., Zambales Prov.</td><td>TNHC62471</td><td>KC822547</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>137</td><td>Kaloula picta</td><td>Philippines, Polillo isl., Quezon Prov.</td><td>USNM-FS56932</td><td>KC822545</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>138</td><td>Kaloula picta</td><td>Philippines, Polillo isl., Quezon Prov.</td><td>USNM-FS-56936</td><td>KC822544</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>139</td><td>Kaloula pulchra</td><td>Thailand, Nong Khai</td><td>KUHE 22206</td><td>AB634632</td><td>AB634690</td><td>Matsui et al. 2011</td></tr><tr><td>140</td><td>Kaloula pulchra</td><td>Thailand, Kanchanaburi, Thong Pha Phum</td><td>KUHE 35171</td><td>AB201183</td><td>AB201194</td><td>Matsui et al. 2005</td></tr><tr><td>141</td><td>Kaloula pulchra</td><td>Myanmar, Sagaing</td><td>USFS34083</td><td></td><td>KC180025</td><td>de Sa et al. 2012</td></tr><tr><td>142</td><td>Kaloula pulchra</td><td>Vietnam, Cat Ba</td><td>ZMMU-A-4761-1</td><td></td><td>PQ268519</td><td>this work</td></tr><tr><td>143</td><td>Kaloula pulchra</td><td>Vietnam, Cat Ba</td><td>ZMMU-A-4761-2</td><td></td><td>PQ268520</td><td>this work</td></tr><tr><td>144</td><td>Kaloula pulchra</td><td>Bangladesh, Sylhet</td><td>IABHU-3781 / Kpul-Bd1</td><td></td><td>AB530543</td><td>Hasan et al. 2012</td></tr><tr><td>145</td><td>Kaloula pulchra</td><td>Bangladesh, Bandarban</td><td>IABHU 3783 / Kpul-Bd2</td><td></td><td>AB530544</td><td>Hasan et al. 2012</td></tr><tr><td>146</td><td>Kaloula pulchra</td><td>Myanmar</td><td>KIZHERP0439</td><td></td><td>JX678910</td><td>Li et al. 2012</td></tr><tr><td>147</td><td>Kaloula pulchra</td><td>Thailand, Ranong</td><td></td><td></td><td>AB530633</td><td>Hasan et al. 2014</td></tr><tr><td>148</td><td>Kaloula pulchra</td><td>Indonesia, Sulawesi, Makassar</td><td></td><td></td><td>AB530639</td><td>Hasan et al. 2014</td></tr><tr><td>149</td><td>Kaloula pulchra</td><td>Malaysia</td><td>VUB0677</td><td></td><td>EF017955</td><td>Van Bocxlaer et al. 2006</td></tr><tr><td>150</td><td>Kaloula pulchra</td><td>–</td><td></td><td>NC006405</td><td></td><td>Zhang et al. 2005</td></tr><tr><td>151</td><td>Kaloula pulchra</td><td>Myanmar</td><td>KIZHERP3003</td><td></td><td>JX678902</td><td>Li et al. 2012</td></tr><tr><td>152</td><td>Kaloula pulchra</td><td>China, Yunnan, Jinghong</td><td>KIZHERP0121</td><td></td><td>JX678903</td><td>Li et al. 2012</td></tr><tr><td>153</td><td>Kaloula pulchra</td><td>China, Yunnan, Mengla</td><td>KIZHERP0129</td><td></td><td>JX678904</td><td>Li et al. 2012</td></tr><tr><td>154</td><td>Kaloula pulchra</td><td>China, Guangxi, Qinzhou</td><td>KIZHERP0171</td><td></td><td>JX678907</td><td>Li et al. 2012</td></tr><tr><td>155</td><td>Kaloula pulchra</td><td>Thailand, Phang Nga</td><td>P994</td><td></td><td>KR827832</td><td>Grosjean et al. 2015</td></tr><tr><td>156</td><td>Kaloula pulchra</td><td>China, Hainan</td><td>GRE119/G027</td><td></td><td>KR827833</td><td>Grosjean et al. 2015</td></tr><tr><td>157</td><td>Kaloula pulchra</td><td>Myanmar, Tanintharyi</td><td>USNM:Herp:586945</td><td></td><td>MG935846</td><td>Mulcahy et al. 2018</td></tr><tr><td>158</td><td>Kaloula pulchra</td><td>Myanmar, Tanintharyi</td><td>USNM:Herp:586946</td><td></td><td>MG935847</td><td>Mulcahy et al. 2018</td></tr><tr><td>159</td><td>Kaloula pulchra</td><td>Myanmar, Sagaing</td><td>USNM:Herp:520322</td><td></td><td>MG935848</td><td>Mulcahy et al. 2018</td></tr><tr><td>160</td><td>Kaloula pulchra</td><td>Myanmar, Sagaing</td><td>USNM:Herp:520326</td><td></td><td>MG935849</td><td>Mulcahy et al. 2018</td></tr><tr><td>161</td><td>Kaloula pulchra</td><td>Myanmar, Bago</td><td>MBM-USNMFS35512</td><td></td><td>MG935850</td><td>Mulcahy et al. 2018</td></tr></tbody></table>
<table><thead><tr><th rowspan="2">No.</th><th>Species</th><th rowspan="2">Locality</th><th rowspan="2">Museum / Sample ID</th><th colspan="2">Accession numbers</th><th rowspan="2">Reference</th></tr><tr><th>Ingroup</th><th>12S rRNA</th><th>16S rRNA</th></tr></thead><tbody><tr><td>162</td><td>Kaloula pulchra</td><td>Myanmar, Yangon</td><td>MBM-JBS19849</td><td></td><td>MG935851</td><td>Mulcahy et al. 2018</td></tr><tr><td>163</td><td>Kaloula pulchra</td><td>Myanmar, Sagaing</td><td>USNM:Herp:523967</td><td></td><td>MG935852</td><td>Mulcahy et al. 2018</td></tr><tr><td>164</td><td>Kaloula pulchra</td><td>Myanmar, Mandalay</td><td>MBM-USNMFS36482</td><td></td><td>MG935853</td><td>Mulcahy et al. 2018</td></tr><tr><td>165</td><td>Kaloula pulchra</td><td>China, Guangxi, Nonggang</td><td>NHMG&lt;CHN&gt;:1106008</td><td></td><td>KC567235</td><td>Mo et al. 2013</td></tr><tr><td>166</td><td>Kaloula pulchra</td><td>China, Guangxi, Nonggang</td><td>NHMG&lt;CHN&gt;:1106009</td><td></td><td>KC567236</td><td>Mo et al. 2013</td></tr><tr><td>167</td><td>Kaloula pulchra</td><td>China</td><td>-</td><td></td><td>AF315162</td><td>Jiang and Zhou, 2001</td></tr><tr><td>168</td><td>Kaloula pulchra</td><td>Southeast Asia</td><td>ZCYK-Kpul1</td><td></td><td>LC640532</td><td>Kambayashi et al. 2022</td></tr><tr><td>169</td><td>Kaloula pulchra</td><td>Bangladesh</td><td>-</td><td></td><td>MN477194</td><td>Rony et al. unpublished</td></tr><tr><td>170</td><td>Kaloula pulchra</td><td>Vietnam, Ha Tinh Prov., Huong Son</td><td>AMCC 106697</td><td>DQ283397</td><td></td><td>Frost et al. 2005</td></tr><tr><td>171</td><td>Kaloula pulchra</td><td>pet trade</td><td>RdS 02</td><td>DQ283398</td><td></td><td>Frost et al. 2005</td></tr><tr><td>172</td><td>Kaloula pulchra</td><td>-</td><td>SIH-09</td><td></td><td>AY330893</td><td>Hoegg et al. 2004</td></tr><tr><td>173</td><td>Kaloula pulchra</td><td>Thailand</td><td>ACD1538</td><td>KC822621</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>174</td><td>Kaloula pulchra</td><td>Laos, Khammouane Prov., Boudalapha</td><td>FMNH225128</td><td>KC822620</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>175</td><td>Kaloula pulchra</td><td>Laos, Bolikhamxay Prov., Thaphabat</td><td>FMNH225129</td><td>KC822619</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>176</td><td>Kaloula pulchra</td><td>Malaysia, Selangor Prov.</td><td>JAM1753</td><td>KC822616</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>177</td><td>Kaloula pulchra</td><td>Malaysia, Pahang Prov., Tioman isl.</td><td>JAM1857</td><td>KC822615</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>178</td><td>Kaloula pulchra</td><td>Malaysia, Pahang, Pulau Tioman</td><td>LSUHC3869</td><td>KC822575</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>179</td><td>Kaloula pulchra</td><td>Malaysia (continental)</td><td>LSUHC3870</td><td>KC852906</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>180</td><td>Kaloula pulchra</td><td>Indonesia, Sumatra isl.</td><td>MF0766</td><td>KC822623</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>181</td><td>Kaloula pulchra</td><td>Indonesia, Sumatra isl.</td><td>MF0812</td><td>KC822624</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>182</td><td>Kaloula pulchra</td><td>China</td><td>NHMS3208</td><td>KC822614</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>183</td><td>Kaloula pulchra</td><td>Indonesia, Sulawesi isl., Propinsi Sulawesi Selatan</td><td>TNHC59422</td><td>KC822618</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>184</td><td>Kaloula pulchra</td><td>Indonesia, Sulawesi isl., Propinsi Sulawesi Selatan</td><td>TNHC59423</td><td>KC822617</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>185</td><td>Kaloula pulchra</td><td>Vietnam</td><td>TZ629</td><td>KC822622</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>186</td><td>Kaloula pulchra</td><td>China</td><td>-</td><td></td><td>AF315130</td><td>Jiang and Zhou, 2001</td></tr><tr><td>187</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Benguet Prov.</td><td>ACD1570</td><td>KC822636</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>188</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Isabela Prov., San Mariano</td><td>ACD1954</td><td>KC822631</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>189</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Isabela Prov., San Mariano</td><td>ACD2043</td><td>KC822632</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>190</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Isabela Prov., San Mariano</td><td>ACD2044</td><td>KC822634</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>191</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Kalinga Prov.</td><td>ACD2032</td><td>KC822633</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>192</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Sierra Madres</td><td>ACD660</td><td>KC822626</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>193</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Kalinga Prov., Balbalan</td><td>ACD756</td><td>KC822628</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>194</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Isabela Prov., San Mariano</td><td>KU326470</td><td>KC822635</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>195</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Cagayan Prov., Pagudpud</td><td>KU328628</td><td>KC822627</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>196</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Cagayan Prov., Gattaran</td><td>RMB4226</td><td>KC822625</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>197</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Kalinga Prov., Balbalan</td><td>TNHC59644</td><td>KC822629</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>198</td><td>Kaloula rigida</td><td>Philippines, Luzon isl., Kalinga Prov., Balbalan</td><td>TNHC60119</td><td>KC822630</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>199</td><td>Kaloula rugifera</td><td>China, Sichuan, Mianyang</td><td>KIZHERP071601</td><td></td><td>JX678911</td><td>Li et al. 2012</td></tr><tr><td>200</td><td>Kaloula rugifera</td><td>China, Sichuan, Mianyang</td><td>KIZHERP071602</td><td></td><td>JX678912</td><td>Li et al. 2012</td></tr><tr><td>201</td><td>Kaloula rugifera</td><td>China, Sichuan, Mianyang</td><td>CIB20110802028</td><td></td><td>KC567237</td><td>Mo et al. 2013</td></tr><tr><td>202</td><td>Kaloula verrucosa</td><td>China, Sichuan, Huidong</td><td>CIB20090349</td><td></td><td>KC567238</td><td>Mo et al. 2013</td></tr><tr><td>203</td><td>Kaloula verrucosa</td><td>China, Sichuan, Huidong</td><td>CIB20090351</td><td></td><td>KC567239</td><td>Mo et al. 2013</td></tr><tr><td>204</td><td>Kaloula verrucosa</td><td>China, Yunnan, Chuxiong</td><td>KIZHERP2015</td><td></td><td>JX678900</td><td>Li et al. 2012</td></tr><tr><td>205</td><td>Kaloula verrucosa</td><td>China, Yunnan, Wuliangshan</td><td>KIZHERP2020</td><td></td><td>JX678901</td><td>Li et al. 2012</td></tr><tr><td>206</td><td>Kaloula verrucosa</td><td>China</td><td>NMNS 3246</td><td>KC822507</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>207</td><td>Kaloula walteri</td><td>Philippines, Mindanao isl., Bukidnon Prov., Malagaylay</td><td>ACD994</td><td>KC822640</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>208</td><td>Kaloula walteri</td><td>Philippines, Polillo isl., Quezon Prov., Polillo</td><td>KU303285</td><td>KC822642</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>209</td><td>Kaloula walteri</td><td>Philippines, Luzon isl., Quezon Prov., Lucban</td><td>KU327338</td><td>KC822639</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>210</td><td>Kaloula walteri</td><td>Philippines, Luzon isl., Quezon Prov., Tayabas</td><td>RMB3701</td><td>KC822641</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>211</td><td>Kaloula walteri</td><td>Philippines, Luzon isl., Quezon Prov., Lucban</td><td>TNHC59667</td><td>KC822637</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>212</td><td>Kaloula walteri</td><td>Philippines, Luzon isl., Quezon Prov., Lucban</td><td>TNHC60116</td><td>KC822638</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>213</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Laguna Prov.</td><td>ACD1692</td><td>KC822602</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>214</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Sierra Madres</td><td>ACD2479</td><td>KC822606</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>215</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Isabela Prov., Palanan</td><td>ACD650</td><td>KC822607</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>216</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Laguna Prov., Los Banos</td><td>ACD943</td><td>KC822597</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>217</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Laguna Prov., Los Banos</td><td>ACD945</td><td>KC822598</td><td></td><td>Blackburn et al. 2013</td></tr></tbody></table>
<table><thead><tr><th rowspan="2">No.</th><th>Species</th><th rowspan="2">Locality</th><th rowspan="2">Museum / Sample ID</th><th colspan="2">Accession numbers</th><th rowspan="2">Reference</th></tr><tr><th>Ingroup</th><th>12S rRNA</th><th>16S rRNA</th></tr></thead><tbody><tr><td>218</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Laguna Prov., Los Banos</td><td>FMNH267555</td><td>KC822599</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>219</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Aurora Prov., Aurora Memorial NP</td><td>RMB750</td><td>KC822601</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>220</td><td>Kaloula sp.</td><td>Philippines, Luzon isl., Aurora Prov., Aurora Memorial NP</td><td>RMB783</td><td>KC822600</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>221</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov., Sibalom</td><td>GVAG253</td><td>KC822515</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>222</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov., Sibalom</td><td>GVAG255</td><td>KC822514</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>223</td><td>Kaloula sp.</td><td>Philippines, Sibuyan isl., Rombolon Prov.</td><td>KU328607</td><td>KC822511</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>224</td><td>Kaloula sp.</td><td>Philippines, Sibuyan isl., Rombolon Prov.</td><td>KU328608</td><td>KC822512</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>225</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov.</td><td>MG012</td><td>KC822513</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>226</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov.</td><td>TNHC56341</td><td>KC822516</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>227</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov.</td><td>TNHC56343</td><td>KC822517</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>228</td><td>Kaloula sp.</td><td>Philippines, Samar isl., Eastern Samar Prov., Taft</td><td>KU310699</td><td>KC822587</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>229</td><td>Kaloula sp.</td><td>Philippines, Leyte isl., Leyte Prov., Danao</td><td>KU328632</td><td>KC822584</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>230</td><td>Kaloula sp.</td><td>Philippines, Leyte isl., Leyte Prov., Danao</td><td>KU328633</td><td>KC822585</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>231</td><td>Kaloula sp.</td><td>Philippines, Leyte isl., Leyte Prov., Baybay</td><td>KU328645</td><td>KC822586</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>232</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov., Sibalom</td><td>MG0000</td><td>KC822611</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>233</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov., Sibalom</td><td>MG0001</td><td>KC822612</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td>234</td><td>Kaloula sp.</td><td>Philippines, Panay isl., Antique Prov., Sibalom</td><td>MG0002</td><td>KC822613</td><td></td><td>Blackburn et al. 2013</td></tr><tr><td></td><td>Outgroups</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>235</td><td>Uperodon taprobanicus</td><td>Bandgladesh, Mymensingh</td><td>Ktap-Bd</td><td>AB530545</td><td></td><td>Hasan et al. 2012</td></tr><tr><td>236</td><td>Uperodon taprobanicus</td><td>Sri Lanka</td><td>KUHE-37252</td><td>AB634633</td><td>AB634691</td><td>Matsui et al. 2011</td></tr><tr><td>237</td><td>Uperodon taprobanicus</td><td>–</td><td>–</td><td>AY948729</td><td></td><td>Roelants et al. 2007</td></tr><tr><td>238</td><td>Kalophrynus interlineatus</td><td>–</td><td>KIZ-HERP-0169</td><td>JX678906</td><td></td><td>Li et al. 2012</td></tr></tbody></table>

These characters were measured in specimens from Khammouane Province, Laos, and Dong Nai and Lam Dong provinces, Vietnam, as well as specimens of *K. indochinensis* from the type locality (Gia Lai Province, central Vietnam). We performed a Principal Component Analysis (PCA) in Statistica v10.0 (StatSoft, Inc. 2011) to examine overall morphological variation among the populations of *K. indochinensis* sensu stricto and the populations of *Kaloula* spp. from central Laos and southern Vietnam. We conducted PCA using SVL and size-corrected values for characters 2–23 (their ratios to SVL), following the methodology of Nishikawa et al. (2007). The first two principal components (i.e., those that explained the highest proportion of variance) were extracted for display with an ordination plot.

Furthermore, we compare external morphological characters based on information taken from the literature (e.g., Chan et al. 2013, 2014; Mo et al. 2013).

## Bioacoustic analysis

Male advertisement calls of *Kaloula* sp. were recorded in Cat Tien National Park, near Nam Cat Tien Village, Tan Phu District, Dong Nai Province, Vietnam, on August 4, 2023 (11.410618°N, 107.419334°E; altitude 119 m a.s.l.); the record was taken at an air temperature of 24.0 °C. Male advertisement calls (described as calls hereafter) were recorded with an Apple iPhone X (iPhone 10); the calls were recorded at a distance of approximately 0.1–0.2 m from the calling males; an HTC-2 Digital LCD Thermometer Hygrometer with an outdoor sensor attached was used to take ambient temperatures and humidity at the calling site after the recording.

The software Raven Pro 1.6 (http://www.birds.cornell.edu/raven) was used to analyze the recorded calls. Audio spectrograms were calculated with a fast Fourier transform (FFT) of 512 points, 90% overlap, and 135 Hz grid spacing using the Hanning window. The terminology of call analysis and description using a call-centered approach (defining uninterrupted units as calls whenever they are separated by long silent intervals) follows Koehler et al. (2017). We examined oscillograms (waveforms) and audio spectrograms of vocalizations and measured the call duration (in milliseconds—ms), intercall interval duration (ms), call repetition rate (calls/s and calls/min), number of notes per call (notes/call), harmonic frequency (kHz), and dominant frequency (kHz). We qualitatively compared the call of *Kaloula* sp. from Cat Tien National Park with the call parameters of *K. indochinensis* sensu stricto from Gia Lai Province, Vietnam, given by Nguyen et al. (2022).

## Results

### Genealogical mitochondrial relationships

The trees recovered by the BI and ML analyses featured essentially similar topologies, with the only differences being the relationships between the higher *K. baleata* clades. Our mtDNA-genealogy confirms the monophyly of the genus *Kaloula* (0.98/96; hereafter node support values are given for BIPP/UFB, respectively) with respect to its sister genus *Uperodon* Duméril & Bibron, 1841 (Fig. 2). Relationships between different groups
one from southern Vietnam (1.0/100) and one from central Laos (1.0/100), the latter being most closely related to *K. indochinensis* from central Vietnam (1.0/100).

*Kaloula baleata* sensu lato, which monophyly is robustly supported only in the ML analysis (0.83/98), features three geographically circumscribed mitochondrial lineages (see Fig. 2): (1) *K. baleata* 1 is distributed in Palawan Island of the Philippines and Sulawesi, Sumba, Bali, central Java, and southern Sumatra Islands of Indonesia; (2) *K. baleata* 2 is distributed in Sarawak and Sabah in Borneo and eastern Peninsular Malaysia, including the island of Tioman, as well as western Java, in sympatry with *K. baleata* 1; (3) *K. baleata* 3 is confirmed from a single locality in Tanintharyi Region, southern Myanmar (see Fig. 1).

## Mitochondrial distances

The uncorrected genetic *p*-distances for the 16S rRNA gene fragment among members of the genus *Kaloula* are available in Table 2. Interspecific distances among *Kaloula* species varied from 1.43% (between *K. verrucosa* and *K. nonggangensis*) to 14.62% (between *K. rugifera* and an undescribed species, *Kaloula* sp. from Panay, Philippines). Intraspecific distances were high in several currently recognized species, including *K. kokacii* (up to 3.39%), but especially in *K. indochinensis* from the *K. baleata* complex, where populations from Khammouane Province, Laos, as well as from Dong Nai and Lam Dong provinces, Vietnam, showed *p*-distances from the closest taxon above 4.45% and 4.33%, respectively.

These 16S distances are much higher than the proposed threshold of 3% for species-level divergence in anurans (Vieites et al. 2009), which was also recently supported to delimit species based on instances of reproductive isolation (Dufresnes et al. 2021). Accordingly, these distances are higher than the *p*-distances observed between some sister species pairs in the *K. baleata* complex, namely 3.0% between *K. baleata* and *K. latidisca*. We also report a high interspecific variation in *K. baleata*, namely 2.72% between *K. baleata* 1 and *K. baleata* 2.

## Morphology

Given the limited number of morphometric characters provided in the original description of *K. indochinensis* in Chan et al. (2013), we did not incorporate their data in our analysis, which further avoids measurer bias. Consequently, the morphological data on *K. indochinensis* relies only on our specimens from near the type locality of this species in Kon Ka Kinh National Park in Gia Lai Province, central Vietnam, as well as in Yok Don National Park in Dak Lak Province, central Vietnam. These specimens were confirmed as *K. indochinensis* by the genetic analysis (Fig. 2). The first two principal components (PCs) of the PCA explain 48.72% (PC1: 28.68%, PC2: 20.04%) of the variation among specimens of *K. indochinensis*, *Kaloula* sp. from central Laos, and *Kaloula* sp. from southern Vietnam (Fig. 3). The specimens of each population/candidate species form distinct groups in the morphospace, without overlapping, and also with respect to sex (Fig. 3).

<figure><img src="image_3.png" /><figcaption>**Figure 3.** Scatterplot of first two PCs in the principal component analysis (PCA) on morphological data from *Kaloula indochinensis*, *Kaloula discordia* sp. nov. from southern Vietnam, and *Kaloula laosensis* sp. nov. from Khammouane, central Laos, overlayed by convex hull polygons; filled circles indicate male specimens, and empty circles indicate female specimens. The morphological data used in the analysis is presented in Table 3.</figcaption></figure>
<table><caption>**Table 2.** Uncorrected average interspecific (below and above diagonal) and intraspecific (on the diagonal) genetic *p*-distances for the 16S rRNA mtDNA gene fragment (in percentage) are given for species of the genus Kaloula.</caption><thead><tr><th></th><th></th><th>Species</th><th>1</th><th>2</th><th>3</th><th>4</th><th>5</th><th>6</th><th>7</th><th>8</th><th>9</th><th>10</th><th>11</th><th>12</th><th>13</th><th>14</th><th>15</th><th>16</th><th>17</th><th>18</th><th>19</th><th>20</th><th>21</th><th>22</th><th>23</th><th>24</th></tr></thead><tbody><tr><td>1</td><td>Kaloula discordia sp. nov.</td><td>0.39</td><td>4.45</td><td>6.17</td><td>5.41</td><td>5.40</td><td>5.47</td><td>6.29</td><td>4.33</td><td>7.00</td><td>6.74</td><td>9.57</td><td>11.17</td><td>8.14</td><td>6.25</td><td>6.98</td><td>7.17</td><td>6.08</td><td>7.45</td><td>7.00</td><td>6.65</td><td>12.77</td><td>9.41</td><td>10.61</td><td>6.54</td></tr><tr><td>2</td><td>Kaloula laosensis sp. nov.</td><td>4.45</td><td>0.22</td><td>4.77</td><td>6.81</td><td>6.59</td><td>7.03</td><td>7.09</td><td>6.28</td><td>7.24</td><td>8.14</td><td>10.74</td><td>12.08</td><td>7.93</td><td>7.65</td><td>8.95</td><td>8.47</td><td>7.47</td><td>6.78</td><td>7.01</td><td>7.89</td><td>14.01</td><td>10.69</td><td>10.64</td><td>8.39</td></tr><tr><td>3</td><td>K. indochinensis</td><td>6.17</td><td>4.77</td><td>0.37</td><td>7.24</td><td>7.25</td><td>6.85</td><td>8.20</td><td>6.26</td><td>7.90</td><td>7.45</td><td>11.87</td><td>12.98</td><td>8.37</td><td>7.92</td><td>8.82</td><td>8.70</td><td>6.86</td><td>6.96</td><td>7.29</td><td>7.84</td><td>15.09</td><td>11.44</td><td>11.07</td><td>7.81</td></tr><tr><td>4</td><td>K. baleata 1</td><td>5.41</td><td>6.81</td><td>7.24</td><td>1.37</td><td>2.72</td><td>2.08</td><td>3.22</td><td>4.79</td><td>6.64</td><td>7.07</td><td>9.72</td><td>10.85</td><td>8.05</td><td>6.90</td><td>6.71</td><td>7.37</td><td>6.24</td><td>8.19</td><td>7.21</td><td>6.72</td><td>12.46</td><td>9.26</td><td>11.25</td><td>7.57</td></tr><tr><td>5</td><td>K. baleata 2</td><td>5.40</td><td>6.59</td><td>7.25</td><td>2.72</td><td>2.56</td><td>2.65</td><td>3.96</td><td>5.36</td><td>6.74</td><td>7.29</td><td>10.26</td><td>11.31</td><td>7.92</td><td>7.35</td><td>7.14</td><td>7.74</td><td>6.76</td><td>8.19</td><td>7.04</td><td>6.82</td><td>12.59</td><td>9.67</td><td>10.59</td><td>7.84</td></tr><tr><td>6</td><td>K. baleata 3</td><td>5.47</td><td>7.03</td><td>6.85</td><td>2.08</td><td>2.65</td><td>0.00</td><td>3.00</td><td>4.98</td><td>6.65</td><td>6.44</td><td>9.10</td><td>10.74</td><td>7.57</td><td>6.86</td><td>6.80</td><td>7.06</td><td>5.99</td><td>7.80</td><td>6.51</td><td>6.26</td><td>13.66</td><td>9.00</td><td>9.95</td><td>7.34</td></tr><tr><td>7</td><td>K. latidisca</td><td>6.29</td><td>7.09</td><td>8.20</td><td>3.22</td><td>3.96</td><td>3.00</td><td>1.45</td><td>6.04</td><td>7.28</td><td>7.47</td><td>9.71</td><td>11.01</td><td>8.89</td><td>7.41</td><td>7.26</td><td>7.74</td><td>7.13</td><td>8.43</td><td>7.82</td><td>7.24</td><td>14.05</td><td>9.41</td><td>11.83</td><td>7.66</td></tr><tr><td>8</td><td>K. mediolineata</td><td>4.33</td><td>6.28</td><td>6.26</td><td>4.79</td><td>5.36</td><td>4.98</td><td>6.04</td><td>5.55</td><td>5.89</td><td>5.43</td><td>8.74</td><td>10.49</td><td>6.12</td><td>5.70</td><td>6.95</td><td>6.05</td><td>5.61</td><td>6.81</td><td>5.46</td><td>5.39</td><td>12.30</td><td>8.49</td><td>10.02</td><td>4.97</td></tr><tr><td>9</td><td>K. borealis</td><td>7.00</td><td>7.24</td><td>7.90</td><td>6.64</td><td>6.74</td><td>6.65</td><td>7.28</td><td>5.89</td><td>0.00</td><td>7.43</td><td>10.12</td><td>11.18</td><td>2.07</td><td>7.07</td><td>7.92</td><td>7.46</td><td>6.44</td><td>2.76</td><td>1.93</td><td>6.78</td><td>13.40</td><td>9.91</td><td>10.11</td><td>7.75</td></tr><tr><td>10</td><td>K. conjuncta</td><td>6.74</td><td>8.14</td><td>7.45</td><td>7.07</td><td>7.29</td><td>6.44</td><td>7.47</td><td>5.43</td><td>7.43</td><td>1.39</td><td>8.37</td><td>9.78</td><td>8.18</td><td>5.08</td><td>8.59</td><td>4.56</td><td>3.53</td><td>7.70</td><td>8.00</td><td>3.56</td><td>11.65</td><td>8.05</td><td>7.06</td><td>2.57</td></tr><tr><td>11</td><td>K. kalingensis</td><td>9.57</td><td>10.74</td><td>11.87</td><td>9.72</td><td>10.26</td><td>9.10</td><td>9.71</td><td>8.74</td><td>10.12</td><td>8.37</td><td>2.07</td><td>6.18</td><td>10.05</td><td>9.39</td><td>11.22</td><td>8.23</td><td>7.55</td><td>9.74</td><td>9.43</td><td>7.52</td><td>10.16</td><td>4.43</td><td>11.20</td><td>7.78</td></tr><tr><td>12</td><td>K. kokacii</td><td>11.17</td><td>12.08</td><td>12.98</td><td>10.85</td><td>11.31</td><td>10.74</td><td>11.01</td><td>10.49</td><td>11.18</td><td>9.78</td><td>6.18</td><td>3.39</td><td>11.41</td><td>10.64</td><td>11.73</td><td>9.29</td><td>9.26</td><td>10.64</td><td>10.51</td><td>8.25</td><td>9.62</td><td>4.48</td><td>12.17</td><td>9.25</td></tr><tr><td>13</td><td>K. monggangensis</td><td>8.14</td><td>7.93</td><td>8.37</td><td>8.05</td><td>7.92</td><td>7.57</td><td>8.89</td><td>6.12</td><td>2.07</td><td>8.18</td><td>10.05</td><td>11.41</td><td>0.00</td><td>7.73</td><td>8.60</td><td>8.77</td><td>8.05</td><td>2.76</td><td>1.43</td><td>8.35</td><td>14.14</td><td>10.02</td><td>11.05</td><td>8.14</td></tr><tr><td>14</td><td>K. pieta</td><td>6.25</td><td>7.65</td><td>7.92</td><td>6.90</td><td>7.35</td><td>6.86</td><td>7.41</td><td>5.70</td><td>7.07</td><td>5.08</td><td>9.39</td><td>10.64</td><td>7.73</td><td>0.31</td><td>9.40</td><td>3.83</td><td>3.15</td><td>7.54</td><td>8.18</td><td>3.51</td><td>11.78</td><td>8.75</td><td>9.18</td><td>4.56</td></tr><tr><td>15</td><td>K. pulchra</td><td>6.98</td><td>8.95</td><td>8.82</td><td>6.71</td><td>7.14</td><td>6.80</td><td>7.26</td><td>6.95</td><td>7.92</td><td>8.59</td><td>11.22</td><td>11.73</td><td>8.60</td><td>9.40</td><td>0.68</td><td>8.88</td><td>8.98</td><td>8.56</td><td>7.93</td><td>9.27</td><td>14.15</td><td>10.63</td><td>12.93</td><td>8.41</td></tr><tr><td>16</td><td>K. rigida</td><td>7.17</td><td>8.47</td><td>8.70</td><td>7.37</td><td>7.74</td><td>7.06</td><td>7.74</td><td>6.05</td><td>7.46</td><td>4.56</td><td>8.23</td><td>9.29</td><td>8.77</td><td>3.83</td><td>8.88</td><td>1.90</td><td>2.73</td><td>8.56</td><td>7.99</td><td>2.45</td><td>9.90</td><td>7.82</td><td>7.74</td><td>3.58</td></tr><tr><td>17</td><td>K. cf. rigida</td><td>6.08</td><td>7.47</td><td>6.86</td><td>6.24</td><td>6.76</td><td>5.99</td><td>7.13</td><td>5.61</td><td>6.44</td><td>3.53</td><td>7.55</td><td>9.26</td><td>8.05</td><td>3.15</td><td>8.98</td><td>2.73</td><td>n/c</td><td>7.59</td><td>7.45</td><td>1.92</td><td>10.72</td><td>7.78</td><td>6.82</td><td>3.88</td></tr><tr><td>18</td><td>K. rugifera</td><td>7.45</td><td>6.78</td><td>6.96</td><td>8.19</td><td>8.19</td><td>7.80</td><td>8.43</td><td>6.81</td><td>2.76</td><td>7.70</td><td>9.74</td><td>10.64</td><td>2.76</td><td>7.54</td><td>8.56</td><td>8.56</td><td>7.59</td><td>0.00</td><td>1.98</td><td>8.12</td><td>14.62</td><td>9.33</td><td>10.70</td><td>7.98</td></tr><tr><td>19</td><td>K. verrucosa</td><td>7.00</td><td>7.01</td><td>7.29</td><td>7.21</td><td>7.04</td><td>6.51</td><td>7.82</td><td>5.46</td><td>1.93</td><td>8.00</td><td>9.43</td><td>10.51</td><td>1.43</td><td>8.18</td><td>7.93</td><td>7.99</td><td>7.45</td><td>1.98</td><td>0.27</td><td>7.48</td><td>13.70</td><td>9.19</td><td>10.44</td><td>7.68</td></tr><tr><td>20</td><td>K. walteri</td><td>6.65</td><td>7.89</td><td>7.84</td><td>6.72</td><td>6.82</td><td>6.26</td><td>7.24</td><td>5.39</td><td>6.78</td><td>3.56</td><td>7.52</td><td>8.25</td><td>8.35</td><td>3.51</td><td>9.27</td><td>2.45</td><td>1.92</td><td>8.12</td><td>7.48</td><td>0.30</td><td>9.38</td><td>6.72</td><td>6.61</td><td>3.19</td></tr><tr><td>21</td><td>Kaloula sp. Panay</td><td>12.77</td><td>14.01</td><td>15.09</td><td>12.46</td><td>12.59</td><td>13.66</td><td>14.05</td><td>12.30</td><td>13.40</td><td>11.65</td><td>10.16</td><td>9.62</td><td>14.14</td><td>11.78</td><td>14.15</td><td>9.90</td><td>10.72</td><td>14.62</td><td>13.70</td><td>9.38</td><td>0.00</td><td>8.51</td><td>12.80</td><td>10.47</td></tr><tr><td>22</td><td>Kaloula sp. East Luzon</td><td>9.41</td><td>10.69</td><td>11.44</td><td>9.26</td><td>9.67</td><td>9.00</td><td>9.41</td><td>8.49</td><td>9.91</td><td>8.05</td><td>4.43</td><td>4.48</td><td>10.02</td><td>8.75</td><td>10.63</td><td>7.82</td><td>7.78</td><td>9.33</td><td>9.19</td><td>6.72</td><td>8.51</td><td>1.85</td><td>10.44</td><td>7.51</td></tr><tr><td>23</td><td>Kaloula sp. Samar & Leyte</td><td>10.61</td><td>10.64</td><td>11.07</td><td>11.25</td><td>10.59</td><td>9.95</td><td>11.83</td><td>10.02</td><td>10.11</td><td>7.06</td><td>11.20</td><td>12.17</td><td>11.05</td><td>9.18</td><td>12.93</td><td>7.74</td><td>6.82</td><td>10.70</td><td>10.44</td><td>6.61</td><td>12.80</td><td>10.44</td><td>1.56</td><td>7.78</td></tr><tr><td>24</td><td>Kaloula sp. Sibuyan</td><td>6.54</td><td>8.39</td><td>7.81</td><td>7.57</td><td>7.84</td><td>7.34</td><td>7.66</td><td>4.97</td><td>7.75</td><td>2.57</td><td>7.78</td><td>9.25</td><td>8.14</td><td>4.56</td><td>8.41</td><td>3.58</td><td>3.88</td><td>7.98</td><td>7.68</td><td>3.19</td><td>10.47</td><td>7.51</td><td>7.78</td><td>1.66</td></tr></tbody></table>
## Bioacoustics

The advertisement call of *Kaloula* sp. from southern Vietnam is documented based on the recordings of one calling male individual. Males of *Kalophrynus interlineatus* (Blyth, 1855), *Occidozyga martensii* (Peters, 1867), and *Microhyla mukhlesuri* Hasan, Islam, Kuramoto, Kurabayashi & Sumida, 2014, were also calling at the same habitat. The calls of *Kaloula* sp. are slow and low-pitched, resonant, booming sounds that resemble the sound of a bicycle horn to the human ear. Call parameters are shown in Table 3, and the sonograms and waveforms of the call are presented in Fig. 4. The call consists of a single note with 11–13 pulses with an average duration of 190.8 ms (144.8–260.6 ms, N = 23). Calls were repeated at a rate of one call per second, 56.5 (42–71) calls per minute, and had an average intercall interval of 1,222.9 ms (327.1–8,954.4 ms, N = 23). The fundamental frequency was not evident, and the average dominant frequency was 0.30–0.45 kHz. Three harmonics were detected at 1.0, 1.2, and 1.5 kHz (Table 3, Fig. 4). The call of the Laos population of *Kaloula* sp. was not recorded.

## Systematics

The two populations of *Kaloula* spp. from Laos and southern Vietnam belong to the *K. baleata* species complex and are characterized by divergent mitochondrial lineages and distinctive morphologies. Their amount of 16S divergence (Table 2), together with the substantial morphological differentiation between them and the closely related species *K. indochinensis* (Fig. 3), suggests a species-level split. Based on our mitochondrial genealogy, the species from Khammouane Province of Laos is the sister species of *K. indochinensis*, while the species from southern Vietnam is the sister species of a clade including *K. indochinensis* and the Laos species (Fig. 2). In addition to diagnostic morphological differences (see below), the three Indochinese species of the *K. baleata* species complex also feature stable differences in coloration, as well as differences in bioacoustic characteristics for the studied Vietnamese species. The distribution of all members of the *K. baleata* species complex in Indochina, following our results, is detailed in Fig. 5. Given the congruence of morphological, bioacoustic, and molecular data in suggesting a deep divergence of the Laos and southern Vietnam *Kaloula* sp. populations from all currently recognized species in the genus, we herein describe them as two new species of *Kaloula*.

### Kaloula discordia Poyarkov, Gorin, Bragin & Nguyen, sp. nov.

https://zoobank.org/3E487D92-32B2-447E-AB70-0059D2F504A9
Figs 2, 4, 6–8, 9C, Tables 3, 4

**Chresonymy.** *Kaloula baleata* [partim]—Orlov et al. (2002: 99); Nguyen et al. (2005: 43); Orlov and Ananjeva (2007: 148); Nguyen et al. (2009: 94).

*Kaloula indochinensis* [partim]—Chan et al. (2013: 334, 2014: 577); Chandramouli and Prasad (2018: 52); Poyarkov et al. (2021b: 39); Vassilieva (2021: 72–73); Holden (2023: 149).

**Holotype.** ZMMU A-8134 (field number NAP-14688), adult male, from a pond in Cat Tien National Park, along the road from park headquarters to the Bau Sau Lake, Dong Nai River valley, Dac Lua Commune, Tan Phu District, Dong Nai Province, Vietnam, collected by N.A. Poyarkov, A.M. Bragin, and V.A. Gorin on 15 June, 2024 (11.44121°N, 107.41312°E; elevation 137 m a.s.l.) (Figs 6, 7C).

**Paratypes (n = 14).** ZMMU A-8135–A-8140 (field numbers NAP-14689–14694), six adult males with collection data same as the holotype; ZMMU A-8141–A-8143 (field numbers NAP-14688–14694), three adult males with collection data same as the holotype; ZISP 15285–15287 (field numbers NAP-14687, NAP-14695–14696), three adult males with collection data same as the holotype; ZMMU A-4739 (field number NAP-01674), adult male from Cat Tien National Park, Dong Nai River valley, Dac Lua Commune, Tan Phu District, Dong Nai Province, Vietnam, collected by N.A. Poyarkov on June 1, 2011 (11.44576°N, 107.38673°E; elevation 128 m a.s.l.); ZMMU A-4642 (field number NAP-00630), adult female from Cat Tien National Park, Dong Nai River valley, Dac Lua Commune, Tan Phu District, Dong Nai Province, Vietnam, collected by N.A. Poyarkov and A.B. Vassilieva on June 2, 2009 (11.44576°N, 107.38673°E; elevation 128 m a.s.l.) (Fig. 8).

**Referred materials (n = 3).** ZMMU-A-4602-1–3 (field numbers NAP-02010, NAP-02086, and NAP-02122), adult female and two subadult females from Cat Tien National Park, Dong Nai River valley, Dac Lua Commune, Tan Phu District, Dong Nai Province, Vietnam, collected by E.A. Galoyan on August 22, 2011, September 15, 2011, and November 11, 2011 (11.419503°N, 107.426442°E; elevation 117 m a.s.l.).

**Diagnosis.** *Kaloula discordia* sp. nov. is distinguished from its congeners by the following combination of morphological characters: (1) medium body size (SVL 42.9–56.2 mm); (2) eyes comparatively small (eye length comprising 60%–75% of snout length); (3) dorsally uniform dark olive-brown; (4) beige-gray ventrally with irregular white mottling on belly and limbs; (5) pale yellow or orange-brown 8-shaped patch on either side of the neck posterior to eyes; (6) yellowish to orange axillary patch present; (7) grayish to beige-yellow inguinal patch present edged with black markings; (8) from gray to beige spot on tibiotarsal articulation present; (9) dark interorbital bar absent; (10) enlarged, widened finger disks (3FDD 6.5%–7.5% of SVL), ca. 1.27 times wider than toe disks; (11) finger subarticular tubercle formula: 1:1:2:2; (12) toe subarticular tubercle formula: 1:1:2:2:2; (13) two metacarpal tubercles not in touch with each other; (14) two metatarsal tubercles, outer metatarsal tubercle rounded, smaller than elongated inner metatarsal tubercle.
<figure><img src="image_4.png" /><figcaption>**Figure 4.** The call of *Kaloula discordia* sp. nov. from Cat Tien NP, Dong Nai Province, Vietnam. **A.** 33 s waveform of relative amplitude (Rel. amp.) and corresponding spectrogram over time; **B.** 1.5 s waveform and corresponding spectrogram of two calls, obtained from the last two calls in (A). Data on acoustic characters is summarized in Table 4.</figcaption></figure>

<table><caption>**Table 3.** Comparison of male advertisement calls for *Kaloula discordia* sp. nov. and other members of the *K. baleata* species complex. NR: not recorded.</caption><thead><tr><th>Species</th><th>Kaloula discordia sp. nov.</th><th>Kaloula indochinensis</th><th>Kaloula baleata</th><th>Kaloula ghoshi</th></tr></thead><tbody><tr><td>Locality</td><td>Cat Tien NP, Tan Phu District, Dong Nai Province, Vietnam</td><td>Kon Von II village, Dak Roong Commune, K'Bang District, Gia Lai Province, Vietnam</td><td>Kinabalu, Sabah, Malaysia</td><td>Hut Bay, Little Andaman Island</td></tr><tr><td>Coordinates</td><td>11.411°N, 107.419°E</td><td>14.543°N, 108.414°E</td><td>NR</td><td>NR</td></tr><tr><td>Number of calls measured</td><td>23</td><td>40</td><td>NR</td><td>NR</td></tr><tr><td>Call duration (ms)</td><td>186.1 (144–214)</td><td>215.6 (194–250)</td><td>280–300</td><td>400 (320–640)</td></tr><tr><td>Intercall interval (ms)</td><td>1539.4 (851–8954)</td><td>789.3 (481–1627)</td><td>NR</td><td>300 (240–530)</td></tr><tr><td>Call repetition rate (call/s)</td><td>1.0 (0.75–1.21)</td><td>1.0 (0.76–1.25)</td><td>NR</td><td>NR</td></tr><tr><td>Call repetition rate (call/minute)</td><td>56.5 (42–71)</td><td>60.1 (45.9–75.2)</td><td>NR</td><td>NR</td></tr><tr><td>Notes/call</td><td>1</td><td>1</td><td>NR</td><td>NR</td></tr><tr><td>Pulse/call</td><td>11–13</td><td>1*</td><td>Described as pulsed</td><td>7–25</td></tr><tr><td>Dominant frequency (kHz)</td><td>0.38 (0.30–0.45)</td><td>0.38 (0.34–0.43)</td><td>1–3</td><td>1</td></tr><tr><td>2<sup>nd</sup> harmonic (kHz)</td><td>1.0</td><td>0.8</td><td>NR</td><td>NR</td></tr><tr><td>3<sup>nd</sup> harmonic (kHz)</td><td>1.2</td><td>1.2</td><td>NR</td><td>NR</td></tr><tr><td>4<sup>nd</sup> harmonic (kHz)</td><td>1.5</td><td>1.37</td><td>NR</td><td>NR</td></tr><tr><td>Temperature (°C)</td><td>24.0</td><td>20.5</td><td>23.0</td><td>29.1</td></tr><tr><td>Source</td><td>this study</td><td>Nguyen et al. (2022)</td><td>Malkmus et al. (2002)</td><td>Chandramouli and Prasad (2018)</td></tr></tbody></table>
<table><caption>**Table 4.** Measurements (in mm) of specimens of *Kaloula discordia* sp. nov., *Kaloula laosensis* sp. nov., and *K. indochinensis*. For character abbreviations, see the Materials and methods section. Abbreviations: F – adult female; M – adult male; SF – subadult female; SM – subadult male.</caption><thead><tr><th>Species</th><th>Sample ID</th><th>Sex</th><th>Status</th><th>SVL</th><th>HL</th><th>SL</th><th>EL</th><th>N-EL</th><th>HW</th><th>IND</th><th>IOD</th><th>UEW</th><th>FLL</th><th>LAL</th><th>HAL</th><th>1FL</th><th>IPTL</th><th>OPTL</th><th>3FDD</th><th>HLL</th><th>TL</th><th>FL</th><th>OMTL</th><th>1TOEL</th><th>4TDD</th></tr></thead><tbody><tr><td>Kaloula discordia sp. nov.</td><td>ZMMU-A-8134</td><td>M</td><td>holotype</td><td>51.5</td><td>11.7</td><td>5.5</td><td>4.1</td><td>3.9</td><td>14.8</td><td>3.2</td><td>5.3</td><td>3.0</td><td>31.9</td><td>26.4</td><td>15.7</td><td>8.1</td><td>3.0</td><td>3.5</td><td>3.6</td><td>60.0</td><td>14.7</td><td>30.2</td><td>2.1</td><td>7.9</td><td>2.3</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-4739</td><td>M</td><td>paratype</td><td>52.0</td><td>11.9</td><td>5.2</td><td>4.0</td><td>3.9</td><td>15.2</td><td>3.2</td><td>5.3</td><td>2.9</td><td>33.4</td><td>26.8</td><td>15.8</td><td>8.2</td><td>3.0</td><td>3.4</td><td>3.8</td><td>62.7</td><td>15.3</td><td>31.4</td><td>2.5</td><td>7.7</td><td>2.9</td></tr><tr><td>K. discordia sp. nov.</td><td>ZISP 15285</td><td>M</td><td>paratype</td><td>48.0</td><td>10.8</td><td>5.1</td><td>3.1</td><td>3.7</td><td>13.7</td><td>3.1</td><td>5.0</td><td>2.8</td><td>31.2</td><td>24.1</td><td>15.0</td><td>7.8</td><td>2.9</td><td>3.3</td><td>3.2</td><td>58.0</td><td>13.4</td><td>28.9</td><td>2.5</td><td>7.9</td><td>2.6</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8135</td><td>M</td><td>paratype</td><td>49.6</td><td>11.3</td><td>5.3</td><td>3.4</td><td>3.8</td><td>14.0</td><td>3.2</td><td>4.9</td><td>2.7</td><td>31.5</td><td>25.3</td><td>15.3</td><td>7.9</td><td>2.6</td><td>3.3</td><td>3.5</td><td>59.0</td><td>14.1</td><td>29.7</td><td>2.4</td><td>7.6</td><td>2.8</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8136</td><td>M</td><td>paratype</td><td>45.4</td><td>10.6</td><td>4.9</td><td>3.3</td><td>3.4</td><td>13.8</td><td>3.0</td><td>4.8</td><td>2.7</td><td>31.4</td><td>24.2</td><td>14.6</td><td>7.8</td><td>2.9</td><td>2.8</td><td>3.3</td><td>57.4</td><td>14.1</td><td>27.9</td><td>2.2</td><td>7.2</td><td>2.4</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8137</td><td>M</td><td>paratype</td><td>42.9</td><td>10.3</td><td>4.9</td><td>3.0</td><td>3.0</td><td>13.7</td><td>3.0</td><td>4.5</td><td>2.6</td><td>30.8</td><td>24.0</td><td>14.9</td><td>7.8</td><td>2.3</td><td>3.1</td><td>3.0</td><td>56.3</td><td>13.1</td><td>28.5</td><td>2.0</td><td>7.1</td><td>2.4</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8138</td><td>M</td><td>paratype</td><td>48.7</td><td>11.1</td><td>5.3</td><td>3.7</td><td>3.5</td><td>14.7</td><td>3.2</td><td>4.9</td><td>2.9</td><td>31.1</td><td>25.0</td><td>15.4</td><td>7.9</td><td>2.4</td><td>3.0</td><td>3.5</td><td>60.6</td><td>14.4</td><td>30.5</td><td>2.2</td><td>7.4</td><td>2.3</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8139</td><td>M</td><td>paratype</td><td>49.5</td><td>11.2</td><td>5.1</td><td>3.1</td><td>3.5</td><td>15.0</td><td>3.1</td><td>4.7</td><td>2.7</td><td>27.9</td><td>23.0</td><td>14.1</td><td>6.8</td><td>2.7</td><td>3.3</td><td>3.5</td><td>57.6</td><td>14.7</td><td>27.4</td><td>1.9</td><td>6.9</td><td>2.8</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8140</td><td>M</td><td>paratype</td><td>47.2</td><td>10.5</td><td>5.0</td><td>3.4</td><td>3.3</td><td>15.1</td><td>3.0</td><td>4.9</td><td>2.7</td><td>27.9</td><td>23.0</td><td>14.1</td><td>6.8</td><td>2.7</td><td>3.3</td><td>3.5</td><td>57.6</td><td>14.7</td><td>27.4</td><td>1.9</td><td>6.9</td><td>2.8</td></tr><tr><td>K. discordia sp. nov.</td><td>ZISP 15286</td><td>M</td><td>paratype</td><td>52.6</td><td>12.2</td><td>5.8</td><td>3.8</td><td>3.8</td><td>15.7</td><td>3.2</td><td>5.6</td><td>2.8</td><td>32.9</td><td>26.3</td><td>15.6</td><td>7.9</td><td>3.2</td><td>3.6</td><td>4.0</td><td>61.9</td><td>15.1</td><td>31.1</td><td>2.3</td><td>8.2</td><td>3.4</td></tr><tr><td>K. discordia sp. nov.</td><td>ZISP 15287</td><td>M</td><td>paratype</td><td>49.1</td><td>10.6</td><td>5.3</td><td>3.6</td><td>3.5</td><td>14.5</td><td>3.1</td><td>5.0</td><td>2.7</td><td>31.2</td><td>24.6</td><td>15.4</td><td>8.1</td><td>2.8</td><td>3.1</td><td>3.4</td><td>60.6</td><td>14.3</td><td>29.7</td><td>2.6</td><td>8.2</td><td>2.6</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8141</td><td>M</td><td>paratype</td><td>50.0</td><td>11.3</td><td>5.5</td><td>3.5</td><td>3.7</td><td>15.3</td><td>3.2</td><td>5.0</td><td>3.0</td><td>31.8</td><td>25.2</td><td>15.2</td><td>7.5</td><td>2.6</td><td>3.0</td><td>3.3</td><td>60.5</td><td>14.3</td><td>31.0</td><td>1.9</td><td>7.4</td><td>2.6</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8142</td><td>M</td><td>paratype</td><td>52.2</td><td>11.7</td><td>5.1</td><td>3.6</td><td>3.2</td><td>15.3</td><td>3.2</td><td>5.0</td><td>2.8</td><td>31.9</td><td>25.1</td><td>15.6</td><td>7.9</td><td>2.5</td><td>3.0</td><td>3.6</td><td>61.0</td><td>14.1</td><td>30.8</td><td>1.9</td><td>7.2</td><td>2.7</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-8143</td><td>M</td><td>paratype</td><td>51.5</td><td>11.7</td><td>5.5</td><td>3.4</td><td>3.6</td><td>15.0</td><td>3.2</td><td>5.1</td><td>2.7</td><td>32.1</td><td>25.4</td><td>15.2</td><td>7.9</td><td>2.9</td><td>3.1</td><td>3.7</td><td>58.5</td><td>13.4</td><td>29.9</td><td>2.4</td><td>7.5</td><td>2.6</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-4642</td><td>F</td><td>paratype</td><td>56.2</td><td>12.8</td><td>6.2</td><td>4.1</td><td>4.2</td><td>17.0</td><td>3.8</td><td>5.9</td><td>3.0</td><td>35.9</td><td>28.2</td><td>16.3</td><td>9.3</td><td>3.2</td><td>3.7</td><td>4.0</td><td>67.4</td><td>16.7</td><td>33.3</td><td>3.0</td><td>9.0</td><td>3.0</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-4602-1</td><td>F</td><td></td><td>56.4</td><td>13.0</td><td>5.4</td><td>4.2</td><td>16.9</td><td>3.7</td><td>6.0</td><td>3.0</td><td>35.8</td><td>28.6</td><td>16.6</td><td>10.0</td><td>3.4</td><td>3.4</td><td>3.8</td><td>67.6</td><td>17.2</td><td>33.7</td><td>3.0</td><td>9.1</td><td>2.9</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-4602-2</td><td>SF</td><td></td><td>25.3</td><td>7.0</td><td>3.1</td><td>2.5</td><td>2.8</td><td>7.5</td><td>1.9</td><td>3.2</td><td>1.5</td><td>15.5</td><td>11.9</td><td>7.0</td><td>3.8</td><td>2.0</td><td>1.8</td><td>1.7</td><td>30.2</td><td>7.4</td><td>13.9</td><td>-</td><td>3.5</td><td>1.5</td></tr><tr><td>K. discordia sp. nov.</td><td>ZMMU-A-4602-3</td><td>SF</td><td></td><td>35.0</td><td>8.3</td><td>3.8</td><td>3.0</td><td>2.8</td><td>10.2</td><td>2.6</td><td>3.6</td><td>1.9</td><td>23.2</td><td>14.5</td><td>10.3</td><td>5.9</td><td>2.2</td><td>2.2</td><td>2.6</td><td>40.2</td><td>9.6</td><td>20.4</td><td>1.4</td><td>4.5</td><td>2.0</td></tr><tr><td>Kaloula laosensis sp. nov.</td><td>ZISP 15284</td><td>M</td><td>holotype</td><td>44.0</td><td>10.6</td><td>4.6</td><td>3.3</td><td>3.2</td><td>12.6</td><td>2.7</td><td>4.3</td><td>2.4</td><td>27.6</td><td>21.7</td><td>13.0</td><td>7.5</td><td>2.7</td><td>2.8</td><td>3.4</td><td>49.3</td><td>11.6</td><td>25.4</td><td>2.0</td><td>7.3</td><td>2.2</td></tr><tr><td>K. laosensis sp. nov.</td><td>ZMMU-A-8144</td><td>F</td><td>paratype</td><td>54.8</td><td>12.4</td><td>5.3</td><td>5.0</td><td>5.2</td><td>16.5</td><td>3.6</td><td>5.7</td><td>3.0</td><td>33.8</td><td>28.3</td><td>17.8</td><td>10.4</td><td>3.2</td><td>3.7</td><td>4.3</td><td>66.9</td><td>16.4</td><td>32.5</td><td>2.8</td><td>8.4</td><td>2.8</td></tr><tr><td>Kaloula indochinensis</td><td>ZMMU-A-8147</td><td>M</td><td></td><td>45.3</td><td>11.0</td><td>4.9</td><td>3.9</td><td>3.3</td><td>14.7</td><td>3.1</td><td>5.0</td><td>2.8</td><td>31.5</td><td>23.7</td><td>13.9</td><td>8.1</td><td>3.1</td><td>3.4</td><td>3.8</td><td>55.2</td><td>14.0</td><td>27.2</td><td>2.19</td><td>7.2</td><td>2.3</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8148</td><td>M</td><td></td><td>50.9</td><td>11.6</td><td>5.3</td><td>3.4</td><td>3.8</td><td>15.1</td><td>3.0</td><td>4.5</td><td>2.5</td><td>32.5</td><td>24.8</td><td>14.6</td><td>7.9</td><td>2.9</td><td>3.1</td><td>3.3</td><td>57.3</td><td>14.5</td><td>27.5</td><td>2.72</td><td>8.1</td><td>2.4</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8149</td><td>M</td><td></td><td>42.3</td><td>9.8</td><td>4.8</td><td>3.4</td><td>3.3</td><td>12.0</td><td>2.9</td><td>4.5</td><td>2.4</td><td>28.1</td><td>20.4</td><td>12.6</td><td>7.8</td><td>2.4</td><td>2.6</td><td>2.9</td><td>47.9</td><td>10.9</td><td>24.9</td><td>2.15</td><td>6.2</td><td>2.2</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8150</td><td>M</td><td></td><td>46.3</td><td>10.7</td><td>4.9</td><td>3.5</td><td>3.5</td><td>13.0</td><td>3.1</td><td>4.5</td><td>2.6</td><td>29.9</td><td>22.4</td><td>13.9</td><td>7.9</td><td>2.6</td><td>3.0</td><td>3.4</td><td>50.7</td><td>12.2</td><td>26.2</td><td>2.59</td><td>7.0</td><td>2.4</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8151</td><td>M</td><td></td><td>48.5</td><td>11.2</td><td>5.2</td><td>3.9</td><td>3.5</td><td>15.7</td><td>3.1</td><td>4.7</td><td>2.7</td><td>31.1</td><td>24.0</td><td>14.6</td><td>8.2</td><td>3.2</td><td>3.3</td><td>3.7</td><td>55.1</td><td>12.9</td><td>27.6</td><td>2.44</td><td>7.4</td><td>2.7</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8152</td><td>M</td><td></td><td>49.1</td><td>11.9</td><td>5.3</td><td>4.0</td><td>3.8</td><td>15.0</td><td>3.3</td><td>5.1</td><td>2.8</td><td>30.6</td><td>24.4</td><td>14.9</td><td>8.7</td><td>3.1</td><td>3.2</td><td>3.6</td><td>55.9</td><td>12.7</td><td>27.5</td><td>2.35</td><td>7.4</td><td>2.3</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8153</td><td>M</td><td></td><td>48.8</td><td>11.8</td><td>4.9</td><td>4.0</td><td>3.6</td><td>15.3</td><td>3.0</td><td>5.0</td><td>2.7</td><td>31.3</td><td>24.3</td><td>14.5</td><td>7.8</td><td>2.9</td><td>3.3</td><td>3.6</td><td>53.0</td><td>12.7</td><td>26.2</td><td>2.80</td><td>6.6</td><td>2.5</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8154</td><td>M</td><td></td><td>49.8</td><td>11.9</td><td>5.5</td><td>3.9</td><td>3.5</td><td>16.3</td><td>3.4</td><td>5.0</td><td>2.7</td><td>32.0</td><td>24.3</td><td>15.1</td><td>8.4</td><td>2.7</td><td>3.1</td><td>3.8</td><td>54.5</td><td>12.9</td><td>27.5</td><td>2.62</td><td>7.4</td><td>2.5</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6316</td><td>M</td><td>topotype</td><td>50.4</td><td>11.7</td><td>5.2</td><td>4.0</td><td>3.7</td><td>14.9</td><td>3.2</td><td>4.9</td><td>2.9</td><td>28.1</td><td>22.8</td><td>14.5</td><td>8.6</td><td>3.2</td><td>3.3</td><td>3.1</td><td>54.7</td><td>12.4</td><td>27.0</td><td>2.75</td><td>6.8</td><td>2.3</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6317</td><td>M</td><td>topotype</td><td>47.3</td><td>10.8</td><td>5.1</td><td>3.8</td><td>3.3</td><td>14.1</td><td>3.0</td><td>4.9</td><td>2.9</td><td>26.7</td><td>22.0</td><td>13.9</td><td>8.0</td><td>2.7</td><td>2.9</td><td>2.8</td><td>52.2</td><td>12.2</td><td>25.7</td><td>2.89</td><td>5.9</td><td>1.8</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6318</td><td>M</td><td>topotype</td><td>51.1</td><td>11.6</td><td>5.4</td><td>3.9</td><td>3.5</td><td>16.2</td><td>3.3</td><td>5.2</td><td>2.7</td><td>31.7</td><td>25.2</td><td>15.5</td><td>8.9</td><td>3.4</td><td>3.4</td><td>3.5</td><td>55.8</td><td>13.0</td><td>27.5</td><td>2.84</td><td>7.1</td><td>2.2</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6319</td><td>M</td><td>topotype</td><td>59.6</td><td>12.7</td><td>5.8</td><td>4.1</td><td>4.0</td><td>17.7</td><td>3.4</td><td>5.8</td><td>3.1</td><td>34.3</td><td>28.6</td><td>18.1</td><td>10.4</td><td>3.9</td><td>4.4</td><td>4.2</td><td>63.9</td><td>15.0</td><td>31.6</td><td>2.79</td><td>8.5</td><td>2.8</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6320</td><td>M</td><td>topotype</td><td>51.3</td><td>12.3</td><td>5.3</td><td>4.0</td><td>3.8</td><td>15.9</td><td>3.4</td><td>5.3</td><td>2.9</td><td>30.5</td><td>24.9</td><td>15.7</td><td>9.2</td><td>3.3</td><td>3.2</td><td>3.4</td><td>3.4</td><td>55.4</td><td>13.1</td><td>27.3</td><td>3.22</td><td>7.0</td><td>2.1</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6321</td><td>M</td><td>topotype</td><td>50.0</td><td>12.3</td><td>5.3</td><td>4.0</td><td>3.7</td><td>16.1</td><td>3.4</td><td>5.4</td><td>3.2</td><td>28.4</td><td>24.0</td><td>14.7</td><td>8.8</td><td>2.9</td><td>3.4</td><td>3.4</td><td>3.4</td><td>55.7</td><td>13.4</td><td>27.5</td><td>3.28</td><td>7.8</td><td>2.4</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6322</td><td>M</td><td>topotype</td><td>50.5</td><td>12.4</td><td>6.0</td><td>4.2</td><td>4.2</td><td>16.5</td><td>3.6</td><td>5.7</td><td>3.0</td><td>30.4</td><td>24.9</td><td>15.7</td><td>9.4</td><td>3.5</td><td>3.5</td><td>3.9</td><td>57.8</td><td>13.9</td><td>28.8</td><td>2.88</td><td>7.8</td><td>2.4</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6323</td><td>M</td><td>topotype</td><td>51.7</td><td>11.4</td><td>5.4</td><td>4.0</td><td>3.7</td><td>15.0</td><td>3.6</td><td>5.9</td><td>3.3</td><td>31.3</td><td>25.3</td><td>15.8</td><td>9.2</td><td>3.6</td><td>3.9</td><td>3.2</td><td>58.0</td><td>14.1</td><td>28.0</td><td>2.85</td><td>7.4</td><td>2.4</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6324</td><td>M</td><td>topotype</td><td>50.2</td><td>12.4</td><td>5.5</td><td>4.0</td><td>3.5</td><td>15.7</td><td>2.9</td><td>5.0</td><td>3.1</td><td>29.1</td><td>23.6</td><td>14.6</td><td>8.2</td><td>3.3</td><td>3.4</td><td>3.2</td><td>55.7</td><td>13.6</td><td>26.5</td><td>2.97</td><td>7.0</td><td>2.2</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6325</td><td>M</td><td>topotype</td><td>48.2</td><td>10.7</td><td>5.1</td><td>3.9</td><td>3.7</td><td>14.3</td><td>3.3</td><td>5.1</td><td>3.0</td><td>28.8</td><td>23.1</td><td>14.4</td><td>7.8</td><td>2.7</td><td>2.9</td><td>3.4</td><td>53.9</td><td>12.4</td><td>26.3</td><td>2.65</td><td>6.7</td><td>2.1</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-6315</td><td>F</td><td>topotype</td><td>55.2</td><td>13.1</td><td>6.2</td><td>4.3</td><td>4.4</td><td>17.3</td><td>3.4</td><td>5.8</td><td>3.1</td><td>32.9</td><td>27.9</td><td>18.0</td><td>9.8</td><td>3.2</td><td>3.8</td><td>4.0</td><td>62.7</td><td>14.8</td><td>31.9</td><td>3.08</td><td>8.4</td><td>2.5</td></tr><tr><td>K. indochinensis</td><td>ZMMU-A-8155</td><td>SF</td><td></td><td>36.1</td><td>8.7</td><td>4.4</td><td>3.4</td><td>2.9</td><td>11.6</td><td>2.6</td><td>3.9</td><td>2.1</td><td>24.8</td><td>18.8</td><td>11.7</td><td>5.9</td><td>2.2</td><td>2.4</td><td>3.0</td><td>41.6</td><td>10.4</td><td>21.3</td><td>1.69</td><td>5.7</td><td>2.1</td></tr></tbody></table>
<figure><img src="image_5.png" /><figcaption>**Figure 5.** Known distribution of the *Kaloula baleata* species complex members in Indochina. Dots within symbols denote type localities. Symbol colors correspond to mitochondrial DNA lineages (Figs 1, 2. Note that the tentative assignment of the populations that were not sequenced is based on geographic proximity and/or examination of morphological or photographic data from these populations). Symbol numbers correspond to locality numbers given in Appendix 1.</figcaption></figure>

**Description of the holotype** (Fig. 6). Adult male in a good state of preservation, habitus robust, head wider than long (HW/HL 1.27), snout projecting beyond lower jaw, gently rounded in lateral view (Fig. 6C); truncated in dorsal view; top of head flat; upper eyelid lacking supra-ciliary tubercles; eye length less than snout length (EL/SL 0.74) and less than interorbital distance (EL/IOD 0.76); pupils round; nostrils rounded, placed more towards the lateral sides of the snout, located closer to tip of snout than to eye, relatively close to each other (IND/IOD 0.60); supratympanic fold flat, glandular, rather thin; tympanum not visible (Fig. 6C); dorsal surfaces of body and limbs with sparse tubercules, getting denser backwards; ventral surfaces of body and limbs with flat tubercules (Fig. 6A, B). Cloacal opening unmodified, directed posteriorly. Forelimbs relatively long, more than a half of hind limb length (FLL/HLL 0.53); hand long, comprising more than a half of lower arm length (HAL/LAL 0.60) and almost half of forelimb length (HAL/FLL 0.49); fingers rather robust, notably flattened in cross section; relative finger lengths: I<II<IV<III; fingers free of webbing; terminal digits flattened forming wide transversely expanded disks (Fig. 6E); finger subarticular tubercles distinct, large and round, finger subarticular tubercle formula 1:1:2:2; inner

metacarpal tubercle elongate; outer metacarpal tubercle triangular-shaped, dilated, bigger than inner metacarpal tubercle (OPTL/IPTL 1.18), two metacarpal tubercles not in touch with each other (Figs 6E, 9C); hindlimbs robust, relatively short, not much longer than body length (HLL/ SVL 1.16); relative toe lengths: I<II<V<III<IV; tarsal fold on inner surface of tarsus absent; tips of all toes widened, forming rounded terminal disks; all toe disks having dorso-terminal grooves; the disk on toe IV the largest (Figs 6D, 9C); toe webbing well developed between all toes, reaching disks at all toes except toe IV; webbing formula 1l-2n1.5-2.5nn1.5-3iv3-1.5v; toe subarticular tubercules distinct, rounded; toe subarticular formula 1:1:2:2:2; two metatarsal tubercles, inner metatarsal tubercle elongated, oval; outer metatarsal tubercle smaller, rounded (Figs 6D, 9C).

**Coloration.** In life, the dorsal surfaces of the head and body olive-brown, dorsal surfaces of the limbs grayish-olive with grayish-white spots (Fig. 6A); flanks of the body and lateral sides of the head grayish olive; orange-brown 8-shaped patches on the neck posterior to eyes; bright-orange axillary patch continuing on elbows (Fig. 7C); grayish-white inguinal patch; small grayish-beige spots near tibiotarsal articulation; ventral surfaces of head and body gray, darker on the throat near the jaw; belly and ventral surfaces of limbs pinkish with weak white mottling (Fig. 6B). Iris golden with brown reticulations ventrally and dorsally from the pupil (Figs 6C, 7C). In preservation after one year of storage in ethanol, dorsal coloration faded to dark gray, light patches became less pronounced, and they faded to light gray or grayish-beige, though the pattern generally remained unchanged.

**Measurements of the holotype (in mm)**: SVL 51.5; HL 11.7; HW 14.8; SL 5.5; EL 4.1; N-EL 3.9; IND 3.2; IOD 5.3; UEW 3.0; FLL 31.9; LAL 26.4; HAL 15.7; HLL 60.0; TL 45.0; FL 30.2; IPTL 3.0; OPTL 3.5; 1FL 8.1; 1TOEL 7.9; OMTL 2.1; 3FDD 3.6; 4TDD 2.3.

**Variation.** Table 4 presents the morphometric variation of the type series. Fig. 8 displays the variation in dorsal coloration of the paratypes. In general, all paratypes agree well with the description of the holotype, differing only in the brightness of light inguinal and axillary markings. In male paratypes ZISP 15285 and ZMMU A-8138, the right foot is poorly developed; in male paratype ZMMU A-8139, the right hand is poorly developed.

**Tadpole morphology.** Vassilieva (2021) provided morphometric data and a detailed description of tadpoles of *Kaloula discordia* sp. nov. (as *K. indochinensis*). Vassilieva (2021) noted that tadpoles of ‘*K. indochinensis*’ from Cat Tien NP, Dong Nai Province, southern Vietnam (corresponding to *Kaloula discordia* sp. nov.) differ from *K. indochinensis* sensu stricto from Kon Ka Kinh NP, Gia Lai Province, and Chu Mon Ray NP, Kon Tum Province, central Vietnam, by having comparatively longer spiracle tubes (markedly longer than the vent tubes), a narrower mouth relative to body width, more developed tail musculature (tail base width more than a third of the body width), and a rather contrasting tail coloration.
<figure><img src="image_6.png" /><figcaption>**Figure 6.** Holotype of *Kaloula discordia* sp. nov. (ZMMU A-8134), adult male. **A.** Dorsal view; **B.** Ventral view; **C.** Head in a lateral view; **D.** Plantar view of left foot; **E.** Volar view of left hand. Photographs by N.A. Poyarkov.</figcaption></figure>

**Distribution and natural history.** *Kaloula discordia* sp. nov. is currently reliably known from Dong Nai, Lam Dong, Tay Ninh, and Binh Phuoc provinces of Southern Vietnam and was also reported from the adjacent Mondulkiri Province of eastern Cambodia; the known distribution of the new species is shown in Fig. 5, and the locality information is detailed in Appendix 1. The single Cambodian record of the new species from Phnom Prich Wildlife Sanctuary in Mondulkiri Province, originally reported as *K. indochinensis* by Chan et al. (2013) and Holden (2023), can be confidently assigned to *Kaloula discordia* sp. nov. based on the external morphology and coloration of the recorded specimen and the geographic proximity of the locality to the population of *Kaloula discordia* sp. nov. in Binh Phuoc Province of southern Vietnam. The new species was recorded mostly from relatively low elevations of ca. 70–300 m a.s.l.; in Bao Loc forestry of Lam Dong

Province, the new species was recorded at elevations up to 550 m a.s.l. *Kaloula discordia* sp. nov. is restricted to lowland and hilly, seasonally dry, semi-deciduous, and evergreen monsoon forests of southern Vietnam (see Vassilieva et al. 2016). Male frogs were recorded calling from small temporary pools and flooded areas from May to November (Fig. 7A, B); the peak of the breeding season coincides with heavy rains in June–August. The new species is semi-fossorial and quite elusive and is usually active only after periods of heavy rains; frogs are aestivating during dry periods in leaf litter, underground borrows, or tree hollows (Vassilieva et al. 2016; Holden 2023). Diet consists of ants and beetles (Vassilieva et al. 2016); the new species is a good climber; males can call when perching a few meters above ground level (Holden 2023). The actual distribution of *K. discordia* sp. nov. is still insufficiently known, but we suppose that this species
<figure><img src="image_7.png" /><figcaption>**Figure 7.** Breeding habitats of *Kaloula discordia* sp. nov. (A, B) and lateral view of holotype of *Kaloula discordia* sp. nov. in situ (ZMMU A-8134) (C). Photographs by A.M. Bragin.</figcaption></figure>

is likely to occur in those provinces of southern Vietnam where forests similar in composition are still preserved, including Dak Nong, Binh Duong, Binh Thuan, and Ba Ria-Vung Tau provinces. Syntopic species of amphibians included *Microhyla butleri* Boulenger, 1900; *M. heymonsi* Vogt, 1911; *M. mukhlesuri*; *Kaloula pulchra*; *Fejervarya limnocharis* (Gravenhorst, 1829); *Occidozyga martensii*; *Polypedates megacephalus* Hallowell, 1861; and *Rhaco-phorus annamensis* Smith, 1924.

**Comparisons.** *Kaloula discordia* sp. nov. most closely resembles other frogs of the *K. baleata* species complex in overall morphology; most specifically, it is similar to *K. indochinensis*, with which it was previously confused. From *K. indochinensis*, the new species can be distinguished by having generally smaller eye size in males (EL mean 3.5 ± 0.3 mm [n = 14] vs. mean 3.9 ± 0.2 mm [n = 18]; EL/SVL 0.071 ± 0.004 vs. 0.079 ± 0.004); smaller finger I in males (1FL mean 7.8 ± 0.4 mm [n = 14] vs. mean 8.5 ± 0.7 mm [n = 18]; 1FL/SVL 0.16 ± 0.01 vs. 0.17 ± 0.007); generally smaller inner palmar tubercules in males (IPTL mean 2.8 ± 0.3 mm [n = 14] vs. mean 3.1 ± 0.4 mm [n = 18]; IPTL/SVL 0.056 ± 0.004 vs. 0.062 ± 0.005); and generally smaller outer palmar tubercules in males (OPTL mean 3.2 ± 0.2 mm [n = 14] mm vs. mean 3.3 ± 0.4 mm [n = 18]; OPTL/SVL 0.064 ± 0.004 vs. 0.067 ± 0.005); generally longer hindlimbs in both sexes (HLL mean 59.6 ± 1.9 mm [n = 14] in males, mean

67.5 ± 0.2 mm [n = 2] in females vs. mean 55.2 ± 3.3 mm [n = 18] in males; 62.7 mm [n = 1] in female; HLL/SVL 1.21 ± 0.04 vs. 1.12 ± 0.03 in both sexes); smaller metatarsal tubercule in males (OMTL mean 2.2 ± 0.3 mm [n = 14] vs. mean 2.7 ± 0.3 mm [n = 18]); by having two metacarpal tubercles, see Fig. 9C (vs. three metacarpal tubercles, see Fig. 9B); by having two subarticular tubercles on toe IV, see Fig. 9C (vs. three well-developed subarticular tubercles, see Fig. 9B); and by having olive dorsal coloration (vs. chocolate-brown to dark grayish-brown).

From *K. baleata* (morphological data taken from Chan et al. 2013), *Kaloula discordia* sp. nov. can be distinguished by having generally smaller eyes in both sexes (EL mean 3.5 ± 0.3 mm [n = 14] in males, mean 4.2 ± 0.01 mm [n = 2] in females vs. mean 4.1 ± 0.4 mm [n = 10] in males, mean 4.5 ± 1.7 mm [n = 2] in females); larger distance between nares in both sexes (IND mean 3.1 ± 0.1 mm [n = 14] in males, mean 3.7 ± 0.1 mm [n = 2] in females vs. mean 2.7 ± 0.4 mm [n = 10] in males, mean 3.1 ± 0.30 mm [n = 2] in females); wider finger III disks in both sexes (3FDD mean 3.5 ± 0.3 mm [n = 14] in males, mean 3.9 ± 0.1 mm [n = 2] in females vs. mean 1.7 ± 0.2 mm [n = 10] in males, mean 2.1 ± 0.6 mm [n = 2] in females); smaller metatarsal tubercule in males (OMTL mean 2.2 ± 0.3 mm [n = 14] vs. mean 3.2 ± 0.2 mm [n = 10]); and by having two subarticular tubercles on toe IV, see Fig. 9C (vs. three well-developed subarticular tubercles).
<figure><img src="image_8.png" /><figcaption>**Figure 8.** Variation in dorsal coloration in paratypes of *Kaloula discordia* sp. nov. Scale bar equals 5 cm. Photographs by N.A. Poyarkov.</figcaption></figure>

From *K. latidisca* (morphological data taken from Chan et al. 2014), the new species can be distinguished by having shorter head in males (HL mean 11.2 ± 0.6 mm [n = 14] vs. mean 14.4 ± 0.8 mm [n = 4]); narrower head in males (HW mean 14.7 ± 0.7 mm (n = 14) vs. mean 18.7 ± 1.1 mm [n = 4]); smaller distance between nares in males (IND mean 3.1 ± 0.1 mm [n = 14] vs. mean 4.0 ± 0.1 mm [n = 4]); smaller distance between eyes in males (IOD mean 5.0 ± 0.2 mm [n = 14] vs. mean 5.7 ± 0.4 mm [n = 4]); smaller eye in males (EL mean 3.5 ± 0.3 mm [n = 14] vs. mean 4.6 ± 0.3 mm [n = 4]); smaller metatarsal tubercle in males (OMTL mean 2.2 ± 0.3 mm [=14] vs. mean 3.4 ± 0.1 mm [n = 4]); and by having two subarticular tubercles on toe IV, see Fig. 9C (vs. three subarticular tubercles).

Furthermore, *Kaloula discordia* sp. nov. differs from *K. aureata* Nutphand, 1989, by having an olive dorsum with no dark reticulations (vs. golden dorsum with dark brown reticulations); from *K. borealis* by having wider finger disks (vs. finger tips slightly dilated but not forming

wide disks); by yellowish blotches on flanks absent (vs. present); and by olive dorsal coloration (vs. gray-brown). The new species is further diagnosed from *K. conjuncta* by the stratified coloration on flanks absent (vs. present) and by having distinct outer metatarsal tubercle (vs. weak or indistinct). *Kaloula discordia* sp. nov. differs from *K. ghoshi* by having axillary and inguinal light spots (vs. absent); tubercles on dorsum and venter (vs. smooth skin or with small granules dorsally); and olive dorsal coloration (vs. orange-brown). The new species further differs from *K. kalingensis* by having tubercles on the dorsum (vs. dorsum smooth); by having a distinct outer metatarsal tubercle (vs. indistinct); yellow or orange axillary and inguinal spots (vs. usually absent or small and red if present); lacking light pericloacal ring (vs. present); from *K. kokacii* by tuberculated dorsum (vs. smooth); axillary and inguinal spots (vs. absent); from *K. mediolineata* by lacking dorso-lateral stripes (vs. present); sacral medial stripe absent (vs. present); and finger disks widened (vs. finger disks slightly
dilated but not forming wide disks). *Kaloula discordia* sp. nov. is distinguished from *K. nonggangensis* by lacking protuberant tubercles on the upper surface of finger tips (vs. present); from *K. picta* by lacking dorsolateral stripes (vs. present); by the stratified coloration on flanks absent (vs. present); and by having wide finger disks (vs. finger disks slightly dilated but not forming wide disks). The new species further differs from *K. pulchra* by the absence of dorsolateral stripes (vs. always present); by having axillary and inguinal spots (vs. absent); from *K. rigida* by having wide finger disks (vs. finger disks slightly dilated but not forming wide disks); by the stratified coloration on flanks absent (vs. present); by the absence of dorsolateral stripes (vs. present); and by having axillary and inguinal spots (vs. absent). *Kaloula discordia* sp. nov. differs from *K. rugifera* by having wide finger disks (vs. slightly expanded small finger disks); by having axillary and inguinal spots (vs. absent); from *K. verrucosa* by having wide finger disks (vs. slightly expanded small finger disks); by having axillary and inguinal spots (vs. absent); from *K. walteri* by having wide finger disks (vs. slightly expanded small finger disks); by the stratified coloration on flanks absent (vs. present); having axillary and inguinal spots (vs. absent); and by having a distinct outer metatarsal tubercle (vs. indistinct or absent). *Kaloula discordia* sp. nov. is geographically isolated from other members of the *K. baleata* species complex and most of the other congeners, except *K. pulchra*, with which it occurs in sympatry everywhere throughout its range, and, possibly, *K. mediolineata*, with which the new species might co-occur in the Tay Ninh, Binh Phuoc, and Ba Ria-Vung Tau provinces of southern Vietnam.

**Acoustic data.** For a comparison of the male advertisement calls of *Kaloula discordia* sp. nov. with *K. indochinensis* sensu stricto, see Table 3; data for *K. indochinensis* are taken from Nguyen et al. (2022). The power call parameters of *Kaloula discordia* sp. nov. and *K. indochinensis* are quite similar, with the dominant frequency of the call being the same in the two species (0.38 kHz, range 0.34–0.43 kHz vs. 0.38 kHz, range 0.30–0.45 kHz, respectively) (Table 3). At the same time, *Kaloula discordia* sp. nov. had a slightly shorter call duration (186.1 ms, range 144–214 ms) than *K. indochinensis* (215.6 ms, range 194–250 ms) and a significantly longer intercall interval: 1539.4 ms (range 851–8954 ms) in *Kaloula discordia* sp. nov. vs. 789.3 ms (range 481–1627 ms) in *K. indochinensis*. These differences have to be taken cautiously as the records of calling of the two species were taken with different ambient temperatures (20.5 °C for *K. indochinensis* and 24.0 °C for the new species), which also could contribute to the observed differences in call parameters between the two species. Nguyen et al. (2022) did not calculate the number of pulses per call in *K. indochinensis* and noted the presence of one pulse per call in this species (Nguyen et al. 2022: Table 2), while we recorded 11–13 pulses per call in *Kaloula discordia* sp. nov. These differences actually result from the different terminology used by Nguyen et al. (2022) and our study, as the waveform oscillograms in Nguyen et al. (2022) clearly show the presence of several pulses in each call.

**Etymology.** The specific epithet “*discordia*” is a noun in apposition, in the nominative case, given in reference to the Roman mythological goddess Discordia. According to the poet Hesiod, this goddess personified not only strife and discord but also competition and labor (Hesiod, Theogony: 20–24, 226–230; see Most 2006). The duality of this name echoes the two aspects of the discovery of the new species. The first aspect is the authors’ hard work and laborious approach in collecting data for the description of the new species. The second challenge pertains to the authors’ internal struggle to choose a politically correct and neutral name for the new species. In modern taxonomy, international teams often face the common challenge of strife and competition; however, this can also lead to overall scientific progress. We recommend “*South Vietnamese Painted Frog*” as the common name in English, “*Yuzhnovietnamskiy Bychiy Uzkorot*” as the common name in Russian, and “*Ênh uong Nam bộ*” as the common name in the Vietnamese language.

**Conservation status.** At present, the new species is reliably known only from four localities in southern Vietnam and a single locality in eastern Cambodia (Fig. 5; Appendix 1). The main threats to this species in Vietnam are habitat loss and degradation. The new species is restricted to the lowland monsoon tropical forests of southern Vietnam; it should be noted that these forests during the last 40 years have been subjected to greater anthropomorphic conversion (including logging, agriculture, road construction, and other human activities) than other areas in Vietnam (e.g., De Koninck 1999; Kuznetsov and Kuznetsova 2011; Lau-rance 2007; Meijer 1973; Meyfroidt and Lambin 2008). The range of the new species covers several nature conservation areas of southern Vietnam and Cambodia, including Cat Tien NP (Dong Nai Biosphere Reserve), Bu Gia Map NP, Lo Go-Xa Mat NP (Vietnam), and Phnom Prich Wildlife Sanctuary (Cambodia). Given the lack of comprehensive studies on the adjacent territories, we suggest *Kaloula discordia* sp. nov. be classified as Data Deficient (DD) according to the IUCN’s Red List categories (IUCN 2019).

**Kaloula laosensis Poyarkov, Orlov, Gorin & Milto, sp. nov.**

https://zoobank.org/BF91FDB-DE05-4518-96E5-E9151DDDF87F

Figs 2, 9A, 10–12, Table 4

**Chresonymy.** *Kaloula baleata* [partim]—Orlov and Ananjeva (2007: 148); Nguyen et al. (2009: 94).

*Kaloula indochinensis* [partim]—Chan et al. (2013: 334, 2014: 577); Teynié et al. (2014: 29); Chandramouli and Prasad (2018: 52); Poyarkov et al. (2021b: 39); Holden (2023: 148).

**Holotype.** ZISP 15284 (field label ZISP 199), adult male, Na Home Village, Bouphala District, near the border of Nakai-Nam Theun National Park, Khammouane Province, Laos, collected by N.L. Orlov, S.N. Nguyen, and K.D. Milto on June 18, 2009 (17.544528°N, 105.695278°E; elevation 174 m a.s.l.).
<figure><img src="image_9.png" /><figcaption>**Figure 9.** Morphological details of feet and hands in the three species of the *Kaloula baleata* species complex of eastern Indochina. **A.** Volar view of the left hand and plantar view of the right foot of *Kaloula laosensis* sp. nov.; **B.** Volar view of the left hand and plantar view of the right foot of *K. indochinensis*; **C.** Volar view of the left hand and plantar view of the right foot of *Kaloula discordia* sp. nov. Scale bar equals 5 mm. Drawings by N.A. Poyarkov.</figcaption></figure>

**Paratype (n = 1).** ZMMU A-8144 (field label ZISP 200), adult female from the same location and with the same collection data as the holotype.

**Diagnosis.** *Kaloula laosensis* sp. nov. is distinguished from its congeners by the following combination of morphological attributes: (1) medium size (SVL ranging 44.0–54.8 mm); (2) eyes comparatively large (eye length comprising 75%–100% of snout length); (3) dark-brown dorsally, with numerous irregular dark blotches forming pericloacal ring; (4) gray or beige ventrally with dense white mottling on belly and limbs; (5) orange triangular patch on either side of the neck posterior to eyes; (6) bright-orange axillary patch present; (7) orange inguinal patch present; (8) bright-orange butterfly-shaped blotch above cloaca present; (9) gray spot on tibiotarsal articulation present; (10) dark interorbital bar present; (11) enlarged, widened finger disks (7.8%–7.9% of SVL), ca. 1.53 times wider than toe disks; (12) finger subarticular tubercle formula: 1:1:2:2; (13) toe subarticular tubercle formula: 1:1:2:2:2; (14) three metacarpal tubercles, median metacarpal tubercle in contact with inner metacarpal tubercle; (15) two metatarsal tubercles, inner metatarsal tubercle ca. three times larger than outer metatarsal tubercle; (16) three small supernumerary tubercles at the basis of toes I, III, IV.

**Description of the holotype** (Fig. 10). Adult male in a good state of preservation, habitus robust; head wider than long (HW/HL 1.19); snout projecting beyond lower jaw, truncated in dorsal and lateral views (Fig. 10C); top of head flat; upper eyelid lacking supraciliary tubercles; eye length less than snout length (EL/SL 0.75) and less than interorbital distance (EL/IOD 0.77); pupils round; nostrils rounded, placed more towards the lateral sides of snout, located closer to tip of snout than to eye, relatively close to each other (IND/IOD 0.63); supratympanic fold flat, tubercular; tympanum not visible (Fig. 10C); dorsal surfaces of body and limbs with sparse tubercules, getting denser backwards; ventral surfaces of body and limbs almost smooth (Fig. 10A, B). Cloacal opening unmodified, directed posteriorly. Forelimbs relatively long, more than a half of hind limb length (FLL/HLL 0.63);

hand long, comprising more than a half of lower arm length (HAL/LAL 0.6) and almost half of forelimb length (HAL/FLL 0.47); fingers rather robust, flattened in cross section; relative finger lengths: I<II<IV<III; no webbing between fingers; terminal digits flattened into very wide transversely expanded T-shaped disks (Fig. 10E); finger subarticular tubercles distinct, protuberant, large and round, finger subarticular formula: 1:1:2:2; metacarpal tubercles three (Figs 9A, 10E), inner metacarpal tubercle elongate, flattened; outer metacarpal tubercle oval, dilated, slightly bigger than inner (OPTL/IPTL 1.05), median metacarpal tubercle the smallest, oval-shaped, contacting both the inner and outer metacarpal tubercles (Figs 9A, 10E); hindlimbs robust, relatively short, not much longer than body length (HLL/SVL 1.12); relative toe lengths: I<II<V<III<IV; tarsal fold on inner surface of tarsus absent; tips of all toes widened, forming terminal oval-shaped disks (Figs 9A, 10D); all toe disks having dorso-terminal grooves; the disk on toe IV the largest; toe webbing well developed between all toes, reaching disks at all toes except toe IV; webbing formula 11.5-2u1.5-2.5III1.5-3IV3-2v; toe subarticular tubercles distinct, rounded, protruding; toe subarticular formula: 1:1:2:2:2; two metatarsal tubercles, inner metatarsal tubercle elongated, oval, shovel-shaped; outer metatarsal tubercle smaller, rounded; three small supernumerary tubercles at the basis of toes I, III, IV (Figs 9A, 10D).

**Coloration.** In life, dorsal surfaces of head and body dark brown with contrasting black spots and blotches; black interorbital bar between the upper eyelids; irregular black blotch on the snout; a series of black blotches forming a L-shaped chevron pattern in the scapular region; elongated black blotches on mid-dorsum, body flanks, and sacral area, forming a black pericloacal ring with irregular borders (Figs 2, 11); dorsal surfaces of limbs grayish brown with dark-brown to black blotches; flanks of body and lateral sides of head grayish brown; on lateral sides of belly getting dark-brown with white spots and blotches; pale orange patches on the neck posterior to eyes; bright reddish-orange axillary patch edged with dark brown; bright reddish-orange inguinal patch edged
<figure><img src="image_10.png" /><figcaption>**Figure 10.** Holotype of *Kaloula laosensis* sp. nov. (ZISP 15284), adult male. **A.** Dorsal view; **B.** Ventral view; **C.** Head in a lateral view; **D.** Plantar view of left foot; **E.** Volar view of left hand. Photographs by V.A. Gorin.</figcaption></figure>

with black; bright reddish-orange patch above cloaca in
the center of dark pericloacal ring (Fig. 2); small gray-
ish-beige spots near tibiotarsal articulation; ventral sur-
faces of head and body gray, darker near the jaw; belly
and ventral sides of limbs densely mottled with white
spots and dots (Fig. 10); iris chocolate brown with copper
sparkles dorsally and ventrally (Fig. 11). In preservation
after 15 years of storage in ethanol, dorsal coloration fad-
ed to dark grayish-brown; dark markings on dorsum are
well discernible; light patches became less pronounced
and faded to yellowish or beige in color, though the col-
oration pattern generally remained unchanged (Fig. 10).

**Measurements of the holotype (in mm):** SVL 44.0;
HL 10.6; HW 12.6; SL 4.6; EL 3.3; N-EL 3.2; IND 2.7;
IOD 4.3; UEW 2.4; FLL 27.6; LAL 21.7; HAL 13.0; HLL
49.3; TL 37.0; FL 25.4; IPTL 2.7; OPTL 2.8; 1FL 7.5;
1TOEL 7.3; OMTL 2.0; 3FDD 3.4; 4TDD 2.2.

**Variation.** Morphometric variation of the type series is presented in Table 4. A dorsal view of a female paratype specimen is presented in Fig. 12. In general, the female paratype specimen agrees well with the description of the holotype, diverging only in body size, which is larger than in the holotype (SVL 54.8 mm). Female paratype ZMMU-A-8144 has comparatively fewer dark markings on the dorsum (Fig. 12), but generally, the coloration pattern is similar to that of the holotype. Holden (2023: 148, fig. 152) published a photo of the new species from Laos as ‘*K. indochinensis*,’ which has bright-red axillary, inguinal, and pericloacal patches and lighter grayish-brown dorsal coloration.

**Tadpole morphology.** Currently, data on the larval
morphology of *Kaloula laosensis* sp. nov. are lacking.

**Distribution and natural history.** *Kaloula laosensis* sp. nov. is currently reliably known only from two localities in the Khammouane Province of Laos; the distribution
<figure><img src="image_11.png" /></figure>

<figure><img src="image_12.png" /><figcaption>**Figure 11.** Dorsolateral (A) and frontal (B) views of the holotype of *Kaloula laosensis* sp. nov. in situ (ZISP 15284, male). Photographs by N.L. Orlov.</figcaption></figure>

of the new species is shown in Fig. 5, and the locality information is detailed in Appendix 1. In Nakai-Nam Theun NP and in Hin Nam No NP, Khammouane Province, Laos, the new species was recorded from limestone evergreen tropical forests at relatively low elevations of ca. 100–300 m asl. The data on the natural history of *Kaloula laosensis* sp. nov. is scarce; it inhabits primarily lowland forests and is quite secretive, emerging only after heavy rains from June to September, when depressions in the forest floor are flooded and form temporary pools. The new species is likely associated with limestone karst landscapes of central Laos; the actual distribution of *Kaloula laosensis* sp. nov. is unknown, but we suppose that this species is likely to occur in adjacent provinces of Vietnam (Ha Tinh and Quang Binh), which harbor limestone forests similar in composition. Details about the new species' reproductive biology and diet are unknown.

**Comparisons.** *Kaloula laosensis* sp. nov. most closely resembles other members of the *K. baleata* species complex in overall morphology, and comparison with these species appears to be the most pertinent. Most specifically, the new species superficially resembles *K. indochinensis*, with which it was previously confused. From *K. indochinensis*, *Kaloula laosensis* sp. nov. can be distinguished by having relatively longer head in males (HL/SVL 0.24 [n = 1] by our data or mean 0.25 ± 0.1 [n = 4] reported by Chan et al. 2013) vs. mean 0.23 ± 0.1 [n = 18] by our data

or [n = 25] reported by Chan et al. 2013); relatively wider disk of finger III in males (3FDD/HAL 0.26 ± 0.1 [n = 1] vs. mean 0.23 ± 0.1 [n = 18]); relatively longer first toe in males (1TOEL/FL 0.29 ± 0.1 [n = 1] vs. mean 0.26 ± 0.1 [n = 18]); by having three metacarpal tubercles with median metacarpal tubercle touching both the inner and outer metacarpal tubercles, see Fig. 9A (vs. median metacarpal tubercle small, not touching the inner metacarpal tubercle, see Fig. 9B); by having two subarticular tubercles on toe IV, see Fig. 9A (vs. three well-developed subarticular tubercles, see Fig. 9B); by having supranumerary tubercles at the basis of toes I, III, and IV, see Fig. 9A (vs. supranumerary tubercles absent, see Fig. 9B); by the presence of numerous contrasting black markings on dorsum (vs. absent or indistinct); and brownish dorsal coloration (vs. chocolate to dark grayish-brown).

From *K. baleata*, *Kaloula laosensis* sp. nov. can be distinguished by having a relatively larger distance between nares in males (IND/HW 0.21 [n = 1] by our data or [n = 6] reported by Chan et al. (2013) vs. 0.18 ± 0.1 [n = 10] reported by Chan et al. (2013)); wider finger III disks in males (3FDD 3.4 ± 0.1 mm [n = 1] by our data) or mean 3.1 ± 0.3 mm [n = 6] reported by Chan et al. (2013) vs. 1.7 ± 0.2 mm [n = 10] reported by Chan et al. (2013)); by having two subarticular tubercles on toe IV, see Fig. 9A (vs. three well-developed tubercles); and by having supranumerary tubercles at the basis of toes I, III, and IV, see Fig. 9A (vs. supranumerary tubercles absent).

From *K. latidisca*, *Kaloula laosensis* sp. nov. can be distinguished by having relatively shorter heads in males (HL/SVL 0.24 [n = 1] by our data or mean 0.25 ± 0.1 [n = 4] reported by Chan et al. (2013) vs. mean 0.27 ± 0.1 [n = 4] reported by Chan et al. (2014)); relatively smaller distance between eyes in males (IOD/HW 0.34 [n = 1] by our data or mean 0.33 ± 0.1 [n = 6] reported by Chan et al. (2013) vs. mean 0.30 ± 0.1 [n = 4] reported by Chan et al. (2014); by the presence of numerous black markings on the dorsum (vs. black markings on dorsum absent or few); and by having two subarticular tubercles on toe IV (vs. three well-developed tubercles).

From *Kaloula discordia* sp. nov. (described above), the new species can be distinguished by having a relatively longer head in males (HL/SVL 0.24 [n = 1] by our data or mean 0.25 ± 0.2 [n = 6] reported by Chan et al. (2013) vs. mean 0.22 ± 0.2 [n = 14]); relatively shorter snout in males (SL/HL 0.43 [n = 1] by our data or mean 0.40 ± 0.1 [n = 6] reported by Chan et al. (2013) vs. mean 0.47 ± 0.1 [n = 14]); relatively narrower head in males (HW/HL 1.19 [n = 1] vs. mean 1.31 ± 0.2 [n = 14]); wider finger III disks (3FDD/HAL 0.26 [n = 1] vs. mean 0.23 ± 0.10 [n = 14]); shorter toe I (1TOEL/FL 0.29 [n = 1] vs. mean 0.25 ± 0.1 [n = 14]); by having three metacarpal tubercles, see Fig. 9A (vs. two metacarpal tubercles, see Fig. 9C); by having supranumerary tubercles at the basis of toes I, III, and IV, see Fig. 9A (vs. supranumerary tubercles absent, see Fig. 9C); by the brown coloration of the dorsum (vs. olive); and by the presence of numerous black markings on the dorsum (vs. black markings on dorsum absent or few).
<figure><img src="image_13.png" /><figcaption>**Figure 12.** Dorsal coloration in a single paratype of *Kaloula laosensis* sp. nov. (ZMMU-A-8144, female) Scale bar equals 5 cm. Photograph by V.A. Gorin.</figcaption></figure>

Furthermore, *Kaloula laosensis* sp. nov. differs from *K. aureata* by having a brownish dorsum with no reticulations (vs. golden dorsum with dark brown reticulations); from *K. borealis* by having wide finger disks (vs. finger tips dilated but not forming wide disks); by the absence of yellowish blotches on flanks (vs. present); an by brownish dorsal coloration (vs. gray-brown). *Kaloula laosensis* sp. nov. further differs from *K. conjuncta* by the stratified coloration on flanks absent (vs. present); by having distinct outer metatarsal tubercle (vs. weak or indistinct); from *K. ghoshi* by the presence of axillary and inguinal spots (vs. axillary and inguinal spots absent); by tuberculated dorsum and venter (vs. smooth or with small flat granules dorsally); by brownish dorsal coloration (vs. orange-brown). The new species differs from *K. kalingensis* in that it has tubercles on the dorsum (vs. dorsum smooth), a distinct outer metatarsal tubercle (vs. indistinct), and yellow or orange axillary and inguinal spots (vs. usually absent or small and red if present). *Kaloula laosensis* sp. nov. further differs from *K. kokacii* by the presence of dorsal tubercles, axillary and inguinal spots, and a light pericloacal ring (vs. absent). The new species differs from *K. mediolineata* by dorsolateral stripes and sacral stripe absent (vs. present); by having enlarged, wide finger disks (vs. finger tips slightly dilated but not forming wide disks); from *K. nonggangensis* by protuberant tubercles on the upper surface of finger tips absent (vs. present); from *K. picta* by the absence of dorsolateral stripes and stratified coloration on flanks (vs. present); by having enlarged, wide finger disks (vs. finger tips slightly dilated but not forming wide disks). The new species can be readily diagnosed from *K. pulchra* by the absence of dorsolateral stripes (vs. present) and by having axillary and inguinal spots (vs. absent). *Kaloula laosensis* sp. nov. further differs from *K. rigida* by having enlarged, wide finger disks (vs. finger tips slightly dilated but not forming wide disks), by the absence of dorsolateral stripes and stratified coloration on flanks (vs. present), and

by having axillary and inguinal light spots (vs. absent). The new species differs from *K. rugifera* by having enlarged, wide finger disks (vs. slightly dilated small finger disks) and by having axillary and inguinal spots (vs. absent) and is further different from *K. verrucosa* by having very wide finger disks (vs. small finger disks) and by having axillary and inguinal spots (vs. absent). The new species differs from *K. walteri* by having very wide finger disks (vs. slightly dilated small finger disks), by the absence of stratified coloration on flanks (vs. presence), by having axillary and inguinal light spots (vs. absent), and by having a distinct outer metatarsal tubercle (vs. indistinct or absent). Except for *K. pulchra*, which can be found in central Laos in syntopy with the new species, *Kaloula laosensis* sp. nov. is geographically separated from most of its relatives.

**Acoustic data.** The male advertisement call of *Kaloula laosensis* sp. nov. has not been recorded, and the bioacoustic data on this species is absent.

**Etymology.** The specific epithet “laosensis” is an adjective in the nominative case, given in reference to the new species’ distribution in central Laos. The name also mirrors the specific epithet of *K. indochinensis*, with which the new species was previously confused. We recommend “Laotian Painted Frog” as the common English name, “Laosskiy bychiy uzkorot” as the common name in Russian, “Ênh uōng Lào” as the common name in Vietnamese, and “Õĩyōng ȳaȳ” (“Ung Yang Lao”) as the common name in Lao languages.

**Conservation status.** At present, the new species is known only from two localities in Khammouane Province of Laos (Fig. 5). The main threats to this species in Laos are habitat loss and degradation due to intensified logging and deforestation, namely Nakai-Nam Theun NP and Hin Nam No NP. We propose that the IUCN’s Red List categories (IUCN 2019) classify *Kaloula laosensis* sp. nov. as Data Deficient (DD) due to the absence of comprehensive studies in the adjacent territories.

**Comments.** We assume, based on the distribution of the new species, that several specimens from the type series of *K. indochinensis* in the original description by Chan et al. (2013), collected in Khammouane Province of Laos and labeled as ‘*K. indochinensis* Laos’ or ‘Laos OTU’ throughout the text, actually represent *Kaloula laosensis* sp. nov. These specimens include six adult males stored in the Field Museum of Natural History (FMNH), Chicago, USA, under voucher numbers FMNH 270360–65. The lack of genetic information on these specimens in the original description of *K. indochinensis* prevented Chan et al. (2013) from recognizing their taxonomic distinctiveness.

## Discussion

Until recently, *K. baleata* sensu lato was reported to inhabit a vast area in Southeast Asia, including territories of Indonesia, Malaysia, India, Thailand, Laos, and Vietnam (Blackburn et al. 2013; Chandramouli and Prasad 2018). This was until Blackburn et al. (2013), who published the first comprehensive phylogeny of the genus, revealed that
K. baleata is a complex of (presumably) cryptic species, which motivated morphological investigations and taxonomic revisions (Chan et al. 2013, 2014). Our discovery of *Kaloula discordia* sp. nov. from Southern Vietnam and *Kaloula laosensis* sp. nov. from Central Laos follows the recent descriptions of two new species from Indochina: *K. indochinensis* (see Chan et al. 2013) and *K. latidisca* (see Chan et al. 2014), as well as the elevation of *K. ghoshi* to full species status (see Chandramouli and Prasad 2018). It is quite remarkable that the populations from southern Vietnam and central Laos, which we herein describe as two new species, were well-known to the authors of previous taxonomic studies of this group (Chan et al. 2013, 2014). In fact, Chan et al. (2013) included six specimens from central Laos (FMNH 270360–65) in their morphological study, assigning them to *K. indochinensis* and including them in the type series of this species, but without assessing their genetic divergence from topotypic specimens of *K. indochinensis* from central Vietnam. Similarly, Chan et al. (2013) did not include specimens of the *K. baleata* complex members from southern Vietnam or adjacent Cambodia in their analysis. However, they included these localities in the proposed distribution range of ‘*K. indochinensis*,’ likely based on photo records, likewise assuming their conspecificity. In essence, our study thus unravelled diversity in the *K. baleata* complex that was “hidden in plain sight” and thereby emphasized the need for careful genetic and phenotypic surveys in presumably widespread species of Southeast Asian amphibians. For example, the common practice of including specimens from different localities in a type series may be misleading. Here, the type locality of *K. indochinensis* described by Chan et al. (2013) from Gia Lai Province of Vietnam and the type locality of *Kaloula laosensis* sp. nov. in Central Laos are located in biogeographically and climatically different parts of Indochina, separated from each other by a straight distance of over 500 km (see Poyarkov et al. 2021b, 2023). Hence, including specimens from both areas in a type series without DNA-barcoding at least some representative specimens from each locality appears an unnecessary gamble.

Different research groups have developed a tradition of providing extensive datasets of morphological characters for Asian anurans (e.g., Poyarkov et al. 2014, 2020b, 2021a; Meegaskumbura et al. 2015; Garg et al. 2019; Biju et al. 2020; Suwannapoom et al. 2018; Brakels et al. 2023). However, some researchers continue to present limited datasets of seemingly handpicked morphological characters. In the description of *K. walteri* by Diesmos et al. (2002), authors provided data on a total of 21 morphometric characters, including complete measurements of fore- and hindlimbs. Mo et al. (2013) reduced the number of studied characters to 14 and lacked data on fingers, toes, and respective disks in their description of *K. nonggangensis*. In the description of *K. indochinensis* by Chan et al. (2013), the number of studied characters further decreased to 11, with no morphometric data on forelimbs and only partial data on hindlimbs. In the following paper, Chan et al. (2014) described *K. latidisca* and provided the same set of 11 characters. Finally, in the most recent

taxonomic paper on the genus *Kaloula* (Chandramouli and Prasad 2018), the authors provided measurements on a set of 17 characters of *K. ghoshi*. None of the respective papers, however, explains or at least mentions the observed reduction in the number of studied morphometric characters. The lack of data on the morphology of these relatively newly described species makes it difficult to compare datasets from different studies and hampers further research into species diversity in *Kaloula* frogs (see the Morphology and Comparisons subsections in the Results section). Since amphibian taxonomy relies heavily on morphological descriptions, we emphasize the utmost importance of providing comprehensive and complete datasets for newly discovered and documented species that can then be re-used in future integrative studies.

Our discovery of two new species of the *K. baleata* complex from Indochina corroborates the idea that many species of Southeast Asian amphibians currently considered to be widely distributed may instead still encompass unrecognized diversity, hidden by a lack of comprehensive field surveys and/or the aforementioned deficiency of comprehensive morphological and genetic data (e.g., Chen et al. 2018; Hasan et al. 2019; Gorin et al. 2020; Suwannapoom et al. 2020; Poyarkov et al. 2020a; Lyu et al. 2023; Liu et al. 2024; Trofimets et al. 2024). Here, the new species of *Kaloula* from the Cat Tien National Park in southern Vietnam appears particularly remarkable. Cat Tien National Park covers one of the few remaining masses of lowland monsoon forests in Vietnam and likely represents one of the best herpetologically studied areas in Vietnam (Yushchenko et al. 2023b). Herpetological exploration of this territory has continued for over 40 years with numerous expeditions and long-term monitoring programs by Vietnamese, Russian, German, USA, British, and Canadian herpetologists, the results of which were summarized in a monograph (Vassilieva et al. 2016). With 45 species of amphibians and 107 species of reptiles recorded in the park, Cat Tien NP is unanimously recognized as an important local center of herpetofaunal diversity (Geissler et al. 2011, 2015; Vassilieva et al. 2013, 2016; Holden and Poyarkov 2021; Yushchenko et al. 2023a,b). The new *Kaloula* species from Cat Tien NP thus illustrates the need for herpetological surveys even in presumably well-known Southeast Asian regions and species.

Moreover, our study provides new information on the evolution, diversity, and distribution of other members of the *K. baleata* species complex, specifically *K. latidisca* and *K. baleata*. Chan et al. (2014) described *K. latidisca* from Kedah, northern Peninsular Malaysia, based on phylogenetic data from Blackburn et al. (2013). In Chan et al.’s (2014) phylogenetic reconstruction, *K. latidisca* is suggested as a sister lineage to *K. baleata* sensu lato, which included samples from Java, Palawan, Borneo, Sulawesi, and southern Peninsular Malaysia. Chan et al. (2014) noted that the populations of southern Peninsular Malaysia and Borneo may not be conspecific with either *K. baleata* or *K. latidisca*, but they refrained from further taxonomic revisions and did not discuss the diversity and extent of distribution of members of the *K. baleata* complex in the
Thai-Malay Peninsula. First, our data showed relatively low, although species-level, sequence differentiation at the 16S rRNA gene between *K. latidisca* and the different lineages of *K. baleata* (p = 3.00%–3.96%; Table 2). Second, the monophyly of *K. baleata* with respect to *K. latidisca* is only moderately supported in our analyses (0.83/98, see Fig. 2). Finally, the mitochondrial distribution of *K. latidisca* and the three lineages of *K. baleata* partially overlap with each other, which requires investigating and potentially updating their respective ranges (see Fig. 1). Specifically, we hypothesize the following.

1. Our mitochondrial data suggests that *K. latidisca* inhabits southern peninsular Thailand, including Satun and Suratthani provinces (see Fig. 1); numerous localities of the *K. baleata* complex from the Isthmus of Kra in the north to the Thai-Malay national border in the south likely belong to this species, which is penetrating to Malaysia only in the state of Kedah (see Fig. 5).

2. The populations of the *K. baleata* complex from Tioman Island in southern Peninsular Malaysia, as well as from the Terengganu and Pahang states in northern Peninsular Malaysia, all belong to the *K. baleata* lineage 1 and thus appear conspecific with the populations from western Java based on their lack of mitochondrial differentiation. It appears that the Titiwangsa Mountain Range separates the *K. baleata* clade 1 and *K. latidisca* ranges in Peninsular Malaysia and southern Thailand.

3. The *K. baleata* population from the Tanintharyi Region of southern Myanmar features distinct mtDNA (lineage 3) that is genealogically closer to the mtDNA lineage(s) found in *K. baleata* sensu stricto (lineage 2) than to *K. latidisca* (Fig. 2). At the same time, this lineage likely extends to the Tenasserim Mountains/Tanintharyi Region north of the Isthmus of Kra (Figs 1, 5). This would imply that *K. latidisca* effectively separates the ranges of the *K. baleata* mtDNA lineages restricted to Sundaland and the Tenasserim Mountains north of the Isthmus of Kra.

These distribution patterns, combined with low genetic divergence in the 16S rRNA gene and unclear morphological differentiation between *K. latidisca* and *K. baleata*, stress the need for further integrative studies to clarify the distribution of these species and the taxonomic relevance of their mitochondrial diversity.

Our new data on the diversity and distribution in the *K. baleata* species complex may have important biogeographic implications. In eastern Indochina, the three *Kaloula* species (*K. indochinensis*, *Kaloula laosensis* sp. nov., and *Kaloula discordia* sp. nov.) group into a clade that is sister to the clade including all other populations of the complex, from the Thai-Malay Peninsula and Southeast Asian islands. The distribution areas of the three Indochinese species correspond well to the main biogeographic regions elaborated by Poyarkov et al. (2021b, 2023). Specifically, the range of *K. indochinensis* is associated with the Central Annamites region, while the newly described *Kaloula*

laosensis* sp. nov. and *Kaloula discordia* sp. nov. are restricted to the Northern Annamites and the Central-South Vietnam Lowlands, respectively. In the *K. baleata*-*K. latidisca* clade, the distribution of *K. baleata* lineage 1 encompasses Wallacea (Sulawesi, Sumba, and Palawan islands), Java, and Sumatra, while *K. baleata* lineage 2 extends over Borneo and eastern Peninsular Malaysia but also on Java (although it may be a result of a human-associated dispersal). At the same time, the range of *K. baleata* lineage 3 is restricted to the North Tenasserim Region, namely north of the Isthmus of Kra, while *K. latidisca* likely inhabits the South Tenasserim south of Kra and the South Thai-Malay- an Lowlands Region. The diversification of these lineages and its association with paleographic events shall greatly benefit from phylogeographic analyses that implement nuclear markers, first to be able to assess their genomic divergence and to test for potential hybridization and introgression in areas of mitochondrial sharing.

## Acknowledgements

The fieldwork in Vietnam was completed within the frameworks and with partial financial support from the research project E-1.2 of the Joint Vietnam-Russia Tropical Science and Technology Research Center for 2024 (Task No. 3). Permission to conduct fieldwork in Vietnam was granted by the Bureau of Forestry, the Ministry of Agriculture and Rural Development of Vietnam, and local administrations. Permission to conduct fieldwork and collect specimens in Cat Tien National Park was granted by Agreement #37/HD on the scientific cooperation between Cat Tien National Park and the Joint Vietnam-Russia Tropical Science and Technology Research Centre. Permissions to conduct fieldwork and collect specimens were granted by the Department of Forestry, Ministry of Agriculture and Rural Development of Vietnam (permit numbers #547/TCLN-BTTN; #432/TCLN-BTTN; #822/TCLN-BTTN; #142/SNgV-VP; #1539/TCLN-DDPH; #1700/UBND.VX); the Forest Protection Departments of the People's Committees of Gia Lai Province (permit numbers #530/UBND-NC; #1951/UBND-NV), Phu Yen Province (permit number #05/UBND-KT); by the Department of Forestry, Ministry of Agriculture and Forestry of Lao PDR (permit number #271/08, form #511); and by the Institute of Animals for Scientific Purpose Development (IAD), Bangkok, Thailand (permit number U1-01205-2558). Specimen collection protocols and animal operations followed the Institutional Ethical Committee of Animal Experimentation of the University of Phayao, Phayao, Thailand (certificate number UP-AE61-01-04-0022). We thank Valentina F. Orlova and Roman A. Nazarov (ZMMU) for permission to examine specimens under their care. We thank the members of MSU HerpLab, including T. Matsukoji, S. S. Idiiatullina, N. S. Kliukin, D. V. Arkhipov, and E. N. Solovyeva, for their support and assistance. NAP and AMB are grateful to Andrei N. Kuznetsov, Svetlana P. Kuznetsova, Hoi Dang Nguyen, and Leonid P. Korzoun for their support and organization.
of fieldwork. We express our sincere gratitude to Peter Mikulíček and two anonymous reviewers for numerous helpful comments and suggestions, which allowed us to improve the previous version of the manuscript.

This work was supported by the Russian Science Foundation to N.A. Poyarkov (Grant No. 22-14-00037, specimen collection and preservation, molecular and morphological analyses, data analyses) and the Thailand Science Research and Innovation Fund and the University of Phayao (Unit of Excellence 2025 on Aquatic animals biodiversity assessment (Phase I)) to C. Suwannapoom.

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## Appendix 1

<table><caption>**Table A1.** List of localities of the *Kaloula baleata* complex members in Indochina appearing on Fig. 5. **Symbols:** (1) = Locality number on Fig. 5; (2) = Verified by morphology data (yes/no); (3) = Verified by molecular data (yes/no).</caption><thead><tr><th>Species</th><th>(1)</th><th>(2)</th><th>(3)</th><th>Location</th><th>Sources</th></tr></thead><tbody><tr><td>K. laosensis sp. nov.</td><td>1</td><td>yes</td><td>yes</td><td>Nakai-Nam Theun NP, Khammouan, Laos (type locality)</td><td>Chan et al. (2013); our data</td></tr><tr><td>K. laosensis sp. nov.</td><td>2</td><td>yes</td><td>no</td><td>Hin Nam No NP, Khammouane, Laos</td><td>Luu et al. (2016)</td></tr><tr><td>K. indochinensis</td><td>3</td><td>yes</td><td>no</td><td>Kiat Ngong, Xepian NBCA, Champasak, Laos</td><td>Chan et al. (2013)</td></tr><tr><td>K. indochinensis</td><td>4</td><td>yes</td><td>no</td><td>Chu Mom Ray NP, Kon Tum, Vietnam</td><td>Jestrzemski et al. (2013)</td></tr><tr><td>K. indochinensis</td><td>5</td><td>yes</td><td>yes</td><td>Kon Chu Rang NR, Gia Lai, Vietnam (type locality)</td><td>Chan et al. (2013); our data</td></tr><tr><td>K. indochinensis</td><td>6</td><td>yes</td><td>yes</td><td>Kon Ka Kinh NP, Gia Lai, Vietnam</td><td>Vassilieva et al. (2016); our data</td></tr><tr><td>K. indochinensis</td><td>7</td><td>yes</td><td>no</td><td>An Khe, Gia Lai, Vietnam</td><td>Chan et al. (2013); our data</td></tr><tr><td>K. indochinensis</td><td>8</td><td>yes</td><td>no</td><td>Song Hinh, Phu Yen, Vietnam</td><td>Do et al. (2017)</td></tr><tr><td>K. indochinensis</td><td>9</td><td>yes</td><td>no</td><td>Tuy Hoa, Phu Yen, Vietnam</td><td>Do et al. (2017)</td></tr><tr><td>K. indochinensis</td><td>10</td><td>yes</td><td>yes</td><td>Yok Don NP, Dak Lak, Vietnam</td><td>our data</td></tr><tr><td>K. discordia sp. nov.</td><td>11</td><td>yes</td><td>yes</td><td>Phnum Prech WS, Mondulkiri, Cambodia</td><td>Chan et al. (2013)</td></tr><tr><td>K. discordia sp. nov.</td><td>12</td><td>yes</td><td>no</td><td>Bu Gia Map NP, Binh Phuoc, Vietnam</td><td>Vassilieva et al. (2016); our data</td></tr><tr><td>K. discordia sp. nov.</td><td>13</td><td>yes</td><td>yes</td><td>Bao Lam, Lam Dong, Vietnam</td><td>Vassilieva et al. (2016); https://www.inaturalist.org/observations/212830145</td></tr><tr><td>K. discordia sp. nov.</td><td>14</td><td>yes</td><td>yes</td><td>Cat Tien NP, Dong Nai, Vietnam (type locality)</td><td>our data; https://www.inaturalist.org/observations/168961087</td></tr><tr><td>K. discordia sp. nov.</td><td>15</td><td>yes</td><td>no</td><td>Lo Go-Xa Mat NP, Tay Ninh, Vietnam</td><td>Vassilieva et al. (2016); our data</td></tr><tr><td>K. baleata Clade 3</td><td>16</td><td>yes</td><td>no</td><td>Thungyai Naresuan East, Umphang, Tak, Thailand</td><td>our data</td></tr><tr><td>K. baleata Clade 3</td><td>17</td><td>yes</td><td>no</td><td>Thong Pha Phum, Kanchanaburi, Thailand</td><td>https://www.inaturalist.org/observations/180645435</td></tr><tr><td>K. baleata Clade 3</td><td>18</td><td>yes</td><td>no</td><td>Suan Phueng, Ratchaburi, Thailand</td><td>our data</td></tr><tr><td>K. baleata Clade 3</td><td>19</td><td>yes</td><td>no</td><td>Kaeng Krachan NP, Phetchaburi, Thailand</td><td>https://www.inaturalist.org/observations/153399059; https://www.inaturalist.org/observations/176926007</td></tr><tr><td>K. baleata Clade 3</td><td>20</td><td>no</td><td>yes</td><td>Yeybu, Tanintharyi, Myanmar</td><td>Zug et al. (2018)</td></tr><tr><td>K. baleata Clade 3</td><td>21</td><td>yes</td><td>no</td><td>Saphan, Prachuap Khiri Khan, Thailand</td><td>https://www.inaturalist.org/observations/218052579</td></tr><tr><td>K. latidisca</td><td>22</td><td>yes</td><td>no</td><td>Saeng, Surathani, Thailand</td><td>Chan-ard et al. (2011); our data</td></tr><tr><td>K. latidisca</td><td>23</td><td>yes</td><td>no</td><td>Khao Sok NP, Surat Thani, Thailand</td><td>https://www.inaturalist.org/observations/70379448; https://www.inaturalist.org/observations/191660698</td></tr><tr><td>K. latidisca</td><td>24</td><td>yes</td><td>yes</td><td>Kapong, Phangnga, Thailand</td><td>https://www.inaturalist.org/observations/161810790</td></tr><tr><td>K. latidisca</td><td>25</td><td>yes</td><td>no</td><td>Khao Lak-Lam Ru NP, Phang Nga, Thailand</td><td>Pauwels et al. (2000); https://www.inaturalist.org/observations/194115334</td></tr><tr><td>K. latidisca</td><td>26</td><td>yes</td><td>no</td><td>Thalang, Phuket, Thailand</td><td>https://www.inaturalist.org/observations/202954715</td></tr><tr><td>K. latidisca</td><td>27</td><td>yes</td><td>no</td><td>Khao Phanom Bencha NP, Krabi, Thailand</td><td>our data</td></tr><tr><td>K. latidisca</td><td>28</td><td>yes</td><td>no</td><td>Ban Na San, Surat Thabi, Thailand</td><td>https://www.inaturalist.org/observations/83178477</td></tr><tr><td>K. latidisca</td><td>29</td><td>yes</td><td>no</td><td>Tha Sala, Nakhon Si Thammarat, Thailand</td><td>https://www.inaturalist.org/observations/190990980</td></tr><tr><td>K. latidisca</td><td>30</td><td>yes</td><td>no</td><td>Khao Luang NP, Nakhon Si Thammarat, Thailand</td><td>Chan-ard et al. (2011); our data</td></tr><tr><td>K. latidisca</td><td>31</td><td>yes</td><td>no</td><td>Khao Pu-Khao Ya NP, Phatthalung, Thailand</td><td>https://www.inaturalist.org/observations/92061585</td></tr><tr><td>K. latidisca</td><td>32</td><td>yes</td><td>no</td><td>Namtok Khao Chong, Trang, Thailand</td><td>Chan-ard et al. (2011); our data</td></tr><tr><td>K. latidisca</td><td>33</td><td>yes</td><td>no</td><td>Yan Ta Khao, Trang, Thailand</td><td>https://www.inaturalist.org/observations/55886801</td></tr><tr><td>K. latidisca</td><td>34</td><td>yes</td><td>yes</td><td>Ton Nga Chang WS, Songkhla, Thailand</td><td>https://www.inaturalist.org/observations/114582513</td></tr><tr><td>K. latidisca</td><td>35</td><td>yes</td><td>yes</td><td>Tha Le Ban NP, Satun, Thialnd</td><td>our data</td></tr><tr><td>K. latidisca</td><td>36</td><td>yes</td><td>no</td><td>Tarutao Island, Satun, Thialnd</td><td>Nidup et al. (2013); our data</td></tr><tr><td>K. latidisca</td><td>37</td><td>yes</td><td>no</td><td>Bang Lang NP, Yala, Thailand</td><td>https://www.inaturalist.org/observations/165320389</td></tr><tr><td>K. latidisca</td><td>38</td><td>yes</td><td>yes</td><td>Gubir, Kedah, Malaysia (type locality)</td><td>Chan et al. (2014)</td></tr><tr><td>K. baleata Clade 1</td><td>39</td><td>yes</td><td>yes</td><td>Kampung Merapunh Lama, Pahang, Malaysia</td><td>Chan et al. (2014)</td></tr><tr><td>K. baleata Clade 1</td><td>40</td><td>yes</td><td>no</td><td>Hulu Terengganu, Terengganu, Malaysia</td><td>Badli-Sham et al. (2023)</td></tr><tr><td>K. baleata Clade 1</td><td>41</td><td>yes</td><td>no</td><td>Kuala Tahan, Pahang, Malaysia</td><td>https://www.inaturalist.org/observations/116005614</td></tr><tr><td>K. baleata Clade 1</td><td>42</td><td>yes</td><td>yes</td><td>Tioman Island, Pahang, Malaysia</td><td>Grismer et al. (2006)</td></tr></tbody></table>