A new species of Leptolalax (Anura: Megophryidae) from northern Vietnam

Abstract

We describe a new, medium-sized Leptolalax species from Vietnam. Leptolalax petrops sp. nov. is distinguished from its congeners by a combination of having a medium-sized body (23.6–27.6 mm in 21 adult males, 30.3–47.0 mm in 17 adult females), immaculate white chest and belly, no distinct black markings on the head, highly tuberculate skin texture, toes lacking webbing and with narrow lateral fringes, and a call consisting of an average of four notes and a dominant frequency of 5.6–6.4 kHz (at 24.5–25.3 °C). Uncorrected sequence divergences between L. petrops sp. nov. and all homologous DNA sequences available for the 16S rRNA gene are >8%.

Full text

544
Accepted by M. Vences: 23 Jan. 2017; published: 16 Mar. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2017 Magnolia Press
Zootaxa 4243 (3): 544
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564
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4243.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:DF89AEF5-CBAB-42A1-84A4-66B1BDD505D6
A new species of Leptolalax (Anura: Megophryidae) from northern Vietnam
JODI J. L. ROWLEY
1,2,6
, VINH Q. DAU
3
, HUY D. HOANG
4
, DUONG T. T. LE
4
,
TIMOTHY P. CUTAJAR
1
& TAO T. NGUYEN
5
1
Australian Museum Research Institute, 1 William St, Sydney NSW 2010, Australia
2
Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney NSW
2052, Australia
3
Hong Duc University, 565 Quang Trung Street-Dong Ve Ward-Thanh Hoa City, Vietnam
4
Faculty of Biology - Biotechnology, Vietnam National University Ho Chi Minh City-University of Science, 227 Nguyen Van Cu, Dis-
trict 5, Ho Chi Minh City, Vietnam
5
Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam
6
Corresponding author. E-mail: Jodi.Rowley@austmus.gov.au
Abstract
We describe a new, medium-sized Leptolalax species from Vietnam. Leptolalax petrops sp. nov. is distinguished from its
congeners by a combination of having a medium-sized body (23.6–27.6 mm in 21 adult males, 30.3–47.0 mm in 17 adult
females), immaculate white chest and belly, no distinct black markings on the head, highly tuberculate skin texture, toes
lacking webbing and with narrow lateral fringes, and a call consisting of an average of four notes and a dominant frequen-
cy of 5.6–6.4 kHz (at 24.5–25.3
°
C). Uncorrected sequence divergences between L. petrops sp. nov. and all homologous
DNA sequences available for the 16S rRNA gene are >8%.
Key words: Acoustics, Anura, L. petrops sp. nov., Southeast Asia, Vietnam
Introduction
The Asian frog genus Leptolalax (Dubois 1983) is a highly morphologically conserved assemblage of small frogs
associated with the forest floor and rocky streams in hilly evergreen forest in Southeast Asia, southern China and
northeastern India (Frost 2016). Their morphological conservatism, small size, often cryptic coloration and
behavior have resulted in a significant underestimation of their true diversity. In the last decade, increased survey
efforts and the incorporation of molecular and acoustic data in delineating species boundaries has led to a rapid
increase in the number of known species in the genus. There are currently 50 species of Leptolalax known, over
half of which have been described in the last five years (Frost 2016).
During recent surveys in Lai Chau and Tuyen Quang Provinces in northern Vietnam, specimens of a medium-
sized species of Leptolalax were collected that did not fit the characteristics of known species. Herein we describe
this species of Leptolalax as new based upon morphological, molecular and bioacoustic data.
Materials and methods
We recorded morphological data from specimens fixed in 10% formalin and then stored in 70% ethanol. Specimens
were deposited at the Australian Museum (AMS) and Vietnam National Museum of Nature (VNMN).
Morphometric data were taken (to the nearest 0.1 mm) with digital calipers. Measurements include snout-vent
length (SVL); head length from tip of snout to rear of jaws (HDL); head width at commissure of jaws (HDW);
snout length from tip of snout to anterior corner of eye (SNT); diameter of exposed portion of eyeball (EYE);
interorbital distance (IOD); horizontal diameter of tympanum (TMP); distance from anterior edge of tympanum to

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A NEW MEGOPHRYID FROG SPECIES FROM VIETNAM
posterior corner of eye (TEY); tibia length with hindlimb flexed (TIB); manus length from tip of third digit to
proximal edge of inner palmar tubercle (ML); and pes length from tip of fourth toe to proximal edge of the inner
metatarsal tubercle (PL). Sex was determined by direct observation of calling in life, and the presence of internal
vocal sac openings and/or gonadal inspection. Mass was recorded in life (to the nearest 0.1 g), using Pesola scales.
Geographic coordinates were obtained using a Garmin GPSMAP 60CSx GPS receiver and recorded in datum
WGS 84. We obtained comparative morphological data from museum specimens of Leptolalax and photographs of
these specimens in life (Appendix I) and from the literature: L. aereus (Rowley et al. 2010a), L. alpinis (Fei et al.
1991, 2009, 2010), L. applebyi (Rowley & Cao 2009), L. arayai (Matsui 1997), L. ardens (Rowley et al. 2016), L.
bidoupensis (Rowley et al. 2011), L. botsfordi (Rowley et al. 2013), L. bourreti (Dubois 1983; Ohler et al. 2011), L.
croceus (Rowley et al. 2010b), L. dringi (Dubois 1987; Inger et al. 1995), L. eos (Ohler et al. 2011), L. firthi
(Rowley et al. 2012), L. fuliginosus (Matsui 2006), L. fritinniens (Dehling & Matsui 2013), L. gracilis (Günther
1872; Inger & Stuebing 2005; Dehling 2012a), L. hamidi (Matsui 1997), L. heteropus (Boulenger 1900), L. isos
(Rowley et al. 2015), L. kalonensis (Rowley et al. 2016a), L. kecil (Matsui et al. 2009), L. khasiorum (Das et al.
2010), L. kajangensis (Grismer et al. 2004), L. lateralis (Anderson 1871; Humtsoe et al. 2008), L. laui (Sung et. al.
2014), L. liui (Fei et al. 1991, 2009, 2010), L. maculosus (Rowley et al. 2016a), L. marmoratus (Matsui et al.
2014a), L. maurus (Inger et al. 1997), L. melanoleucus (Matsui 2006), L. melicus (Rowley et al. 2010c), L.
minimus (Ohler et al. 2011), L. nahangensis (Lathrop et al. 1998), L. nokrekensis (Mathew & Sen 2010), L. nyx
(Ohler et al. 2011), L. oshanensis (Liu 1950; Fei et al. 2009, 2010), L. pallidus (Rowley et al. 2016a), L.
pelodytoides (Boulenger 1893, 1908; Ohler et al. 2011), L. pyrrhops (Poyarkov et al. 2015) L. pictus (Malkmus
1992; Malkmus et al. 2002), L. platycephalus (Dehling 2012b), L. pluvialis (Ohler et al. 2000, 2011), L.
sabahmontanus (Matsui et al. 2014b), L. solus (Matsui 2006), L. sungi (Lathrop et al. 1998), L. tadungensis
(Rowley et al. 2016a), L. tamdil (Sengupta et al. 2010), L. tengchongensis Yang et al. 2016), L. tuberosus (Inger et
al.1999; Rowley et al. 2010b), L. ventripunctatus (Fei et al. 1991, 2009, 2010), L. zhangyangpingi (Jiang et al.
2013). Due to the degree of undiagnosed diversity within the genus, where available, we relied on examination of
topotypic material and/or original species descriptions.
Advertisement calls were recorded with an Edirol R-09HR WAVE/MP3 Recorder (96 kHz sampling rate and
24-bit encoding) with a Røde NTG-2 condenser shotgun microphone. Calls were recorded at a distance of
approximately 0.1–0.3 m and ambient air temperatures at calling sites were taken immediately after recordings
using a Kestrel 3500 hand-held weather meter. Calls were analysed with Raven Pro 1.3 software (http://
www.birds.cornell.edu/raven). Audiospectrograms for analysis were calculated with fast-Fourier transform (FFT)
of 512 points, 50% overlap and 172 Hz grid-spacing, using Hanning windows, for figures we used 512 points. In
describing the advertisement calls, we use the definitions of Duellman (1970), except that we define each call as a
vocalisation produced during a single expiration. Temporal and spectral parameters of calls were measured using
the definitions of Cocroft & Ryan (1995), except for fundamental frequency, where the definition of Duellman
(1970) was used. For each call recording, we measured the call duration (ms), call repetition rate (calls/s), intercall
interval (ms), number of notes per call, note duration (ms) and dominant frequency (kHz). Comparative
advertisement call characters for Leptolalax species were taken from references, with advertisement calls known
for 32 of the 50 known species of Leptolalax (Matsui 1997, 2006; Jiang et al. 2002; Malkmus et al. 2002; Matsui et
al. 2009, 2014a, 2014b; Xu et al. 2005; Rowley & Cao 2009; Rowley et al. 2010a, 2010b, 2010c, 2011, 2012,
2013, 2016a, 2016b; Sukumaran et al. 2010; Dehling & Matsui 2013; Poyarkov et al. 2015). To maintain
consistency and facilitate meaningful comparisons, we have used the terminology defined above to compare calls,
regardless of terms used in these references.
Total genomic DNA was extracted from tissues using DNeasy tissue extraction kits (Qiagen). We used the
primers 16SAR and 16SBR of Palumbi et al. (1991) to amplify ~550 base pairs of the 16S rRNA gene for the new
species. Standard PCR protocols were used and PCR products were purified using ExoSap-IT (USB Corporation,
OH, USA). Purified templates were sequenced directly by Macrogen (Seoul, Korea).
In addition to the newly collected specimens, all species belonging to the genus Leptolalax for which
homologous sequences were available (33 of the 50 species) were included in the phylogenetic analysis.
Leptobrachium cf. chapaense and Xenophrys major were included in the analysis as an outgroup. Locality
information and accession numbers for all sequences included in the analysis can be found in Table 1.
Sequences were validated using Sequencher 4.10 (Gene Codes, Ann Arbor, MI) and aligned using MAFFT
(Katoh et al. 2002) on the CIPRES Science Gateway (Miller et al. 2010). We used the Akaike Information

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Criterion as implemented in jModelTest 2.1.6 (Darriba et al. 2012) on the CIPRES Science Gateway (Miller et al.
2010) to select the best-fit model of nucleotide substitution, which was then used in model-based phylogenetic
inference.
TABLE 1. DNA sequences of the mitochondrial 16S rRNA gene used in this study. *generated as part of this study.
Species Locality Voucher no. GenBank no.
Leptolalax petrops sp. nov. Vietnam, Tuyen Quang Province AMS R184826, VNMN 2016
A.06
KY459997*,KY4
59998*
Leptolalax petrops sp. nov. Vietnam, Lai Chau Province AMS R 184845, VNMN 2016
A.16
KY459999*,
KY460000*
Leptolalax aereus Vietnam, Quang Binh Province RH60165 JN848437
Leptolalax applebyi Vietnam, Quang Nam Province AMS R171703 HM133597
Leptolalax arayai Malaysia, Borneo BORNEENSIS 22931 AB847558
Leptolalax ardens Vietnam, Gia Lai Province VNMN 04707/AMS R176454 KR018109
Leptolalax bidoupensis Vietnam, Lam Dong Province AMS R173134 HQ902881
Leptolalax bourreti Vietnam, Lao Cai Province AMS R177673 KR018124
Leptolalax dringi Malaysia, Borneo KUHE:55610 AB847553
Leptolalax eos Laos, Phongsaly Province MNHN:2004.0278 JN848450
Leptolalax firthi Vietnam, Kon Tum Province AMS R176524 JQ739206
Leptolalax fritinniens Malaysia, Borneo KUHE 55371 AB847557
Leptolalax gracilus Malaysia, Borneo KUHE 55624 AB847560
Leptolalax hamidi Malaysia, Borneo KUHE 17545 AB969286
Leptolalax heteropus Malaysia, Peninsular KUHE 15487 AB530453
Leptolalax isos Vietnam, Gia Lai Province VNMN A 2015.4/AMS R176480 KT824769
Leptolalax kalonensis Vietnam, Binh Thuan Province IEBR A.2014.15 KR018114
Leptolalax laui China, Shenzhen SYS A002057 KM014546
Leptolalax liui China, Jiangxi Province SYS A001620 KM014549
Leptolalax maculosus Vietnam, Ninh Thuận Province ZFMK 96600 KR018120
Leptolalax marmoratus Malaysia, Borneo KUHE 53227 AB969289
Leptolalax maurus Malaysia, Borneo SP 21450 AB847559
Leptolalax melicus Cambodia, Ratanakiri Province MVZ 258198 HM133600
Leptolalax minimus Thailand, Chiang Mai Province - JN848369
Leptolalax nyx Vietnam, Ha Giang Province AMNH A163810 DQ283381
Leptolalax oshanensis China, Sichuan Province SYS A001830 KM014810
Leptolalax pallidus Vietnam, Lam Dong Province UNS00510 KR018112
Leptolalax pictus Malaysia, Borneo UNIMAS 8705 KJ831295
Leptolalax pluvialis Vietnam, Lao Cai Province MNHN:1999.5675 JN848391
Leptolalax pyrrhops Vietnam, Lam Dong Province ZMMU A-5208 KP017575
Leptolalax sabahmontanus Malaysia, Borneo BORNEENSIS 12632 AB847551
Leptolalax tadungensis Vietnam, Dak Nong Province UNS00515 KR018121
Leptolalax tengchongensis China, Yunnan Province SYS a004600 KU589210
Leptolalax ventripunctatus Laos, Phongsaly Province MNHN 2005.0116 JN848410
Leptolalax zhangyangpingi Thailand, Chiang Mai Province - JX069979
Leptobrachium cf. chapaense Vietnam, Lao Cai Province AMS R171623 KR018126
Xenophrys major Vietnam, Kon Tum Province AMS R173870 KY476333*

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Bayesian phylogenetic analyses were performed in MrBayes 3.2 (Ronquist et al. 2012) under a GTR+I +Γ
(general time reversible model with Gamma distributed substitution rates and a proportion of the sites invariable;
model parameters estimated during the search). Four independent Markov Chain Monte Carlo searches were run
for one million generations, sampled every 1000 generations, each with four chains, and default priors. Output files
were examined visually in Tracer v1.5 (Rambaut & Drummond 2007), to assess stationarity and determine the
number of generations to remove as burn-in. To be conservative, we considered 100,000 generations from each run
as burn-in, and removed 100 trees before summarizing topology and posterior probabilities. Trees were visualized
using the FigTree v1.4.2 program, available at http://tree.bio.ed.ac.uk/software/figtree/. We consider branches
receiving ≥0.95 posterior probabilities to be well-supported (Wilcox et al. 2002).
Maximum likelihood analyses were performed on the data matrix using RaxML accessed via the CIPRES
Science Gateway (Miller et al. 2009) (www.phylo.org/portal2/) using default settings.
FIGURE 1. Type localities of Leptolalax petrops sp. nov.
Results
The newly collected Leptolalax specimens were recovered within the clade of Leptolalax in the subgenus Lalos
(Delorme et al. 2006) known from northern and central Vietnam, Laos and China: L. aereus, L. bourreti, L. eos, L.
firthi, L. isos, L. laui, L. liui, L. nyx, L. oshanensis, L. pluvalis and L. tengchongensis (Fig. 2). The clade was well-
supported in ML but not Bayesian analysis. Uncorrected 16S rRNA sequence divergences between the newly
collected specimens and all homologous sequences available on GenBank (sequences assigned to 33 species; Table
2) were ≥8.0%. This degree of pairwise divergence in the 16S rRNA gene is much greater than that usually
representing differentiation at the species level in frogs (Vences et al. 2005). The newly collected specimens
clustered together in a well-supported clade. Uncorrected sequence divergence within the two specimens sampled
at each site was 0.0–0.2% and between the two sites (a distance of >180km) was 2.7–3.1% at the 16S gene
fragment analysed.

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    
 Leptolalax aereus 
 Leptolalax applebyi 
 Leptolalax arayai 
 Leptolalax ardens 
 Leptolalax bidoupensis 
 Leptolalax bourreti 
 Leptolalax dringi 
 Leptolalax eos 
 Leptolalax firthi                  
 Leptolalax fritinniens                  
 Leptolalax gracilus                  
 Leptolalax hamidi                  
 Leptolalax heteropus                  
 Leptolalax isos                  
 Leptolalax kalonensis                  
 Leptolalax laui                  
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Zootaxa 4243 (3) © 2017 Magnolia Press
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A NEW MEGOPHRYID FROG SPECIES FROM VIETNAM
7$%/(&RQWLQXHG 
                  
 Leptolalax liui                  
 Leptolalax maculosus                  
 Leptolalax marmoratus                  
 Leptolalax maurus                  
 Leptolalax melicus                  
 Leptolalax minimus                  
 Leptolalax nyx                  
 Leptolalax oshanensis                  
 Leptolalax pallidus                  
 Leptolalax pictus                  
 Leptolalax pluvialis                  
 Leptolalax pyrrhops                  
 Leptolalax sabahmontanus                  
 Leptolalax tadungensis                  
 Leptolalax tengchongensis                  
 Leptolalax ventripunctatus                  
 Leptolalax zhangyangpingi                  
 Leptobrachium FI chapaense                  
 Xenophrys major                  
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7$%/(  FRQW 8QFRUUHFWHG 6 SDL UZLVHG LVWDQFHV DPRQJ QHZO\ FROOHFWHG Leptolalax VSHFLPHQVIURP QRUWKHUQ 9LHWQDPDQG DOO Leptolalax VSHFLHVIRU ZKLFK FRPSDUDEOHVHTXHQFHV DUH DYDLODEOH SOXV
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                     
 Leptolalax kalonensis 
 Leptolalax laui 
 Leptolalax liui  
 Leptolalax maculosus 
 Leptolalax marmoratus 
 Leptolalax maurus                    
 Leptolalax melicus                    
 Leptolalax minimus                    
 Leptolalax nyx                    
 Leptolalax oshanensis                    
 Leptolalax pallidus                    
 Leptolalax pictus                    
 Leptolalax pluvialis                    
 Leptolalax pyrrhops                    
 Leptolalax sabahmontanus                    
 Leptolalax tadungensis                    
 Leptolalax tengchongensis                    
 Leptolalax ventripunctatus                    
 Leptolalax zhangyangpingi                    
 Leptobrachium FI chapaense                    
 Xenophrys major                    


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FIGURE 2. Bayesian inference tree on ~570 bp fragment of the 16S gene for Leptolalax petrops sp. nov., along with
representatives of all Leptolalax species for which comparable sequences are available. Node support is indicated on branches
as Bayesian posterior probabilities (upper half; >0.95= grey, 1 = black) and Maximum Likelihood support (lower; >70%, 100 =
black).
Leptolalax petrops sp. nov.
Figs. 3–4.
Holotype: VNMN 2016 A.06, adult male, 0.5 m from 5 m wide, rocky stream in evergreen forest at Cham Chu
Nature Reserve, Tuyen Quang Province, Vietnam (22.2045
°
N, 105.0754
°
E, 715 m; Fig. 1). Collected on 25 May
2013 by Jodi J. L. Rowley, Dau Quang Vinh, Hoang Duc Huy, and Hoang Anh Tuan.
Paratypes: AMS R 84828, adult female, and AMS R184828, VNMN 2016 A.08, VNMN 2016 A.09, VNMN
2016 A.10, VNMN 2016 A.11, AMS R184829, six adult males, collected at same location and date as holotype;
AMS R184826 and AMS R1848287, two adult males, collected at same location as holotype on 24 May 2013.
AMS R184830, AMS R184832, AMS R184833, three adult males collected at same location as holotype on 26
May 2013. AMS R184831, adult male, collected 2 m from 5 m wide rocky stream in evergreen forest at Cham Chu
Nature Reserve, Tuyen Quang Province, Vietnam (22.2061
°
N, 105.0747
°
E, 587 m) on 26 May 2013. AMS
R184834, adult male collected 0.5 m from 15 m wide, rocky stream in disturbed evergreen forest at Cham Chu
Nature Reserve, Tuyen Quang Province, Vietnam (22.2105
°
N, 105.0619
°
E, 234 m) on 1 June 2013. VNMN 2016
A.12, adult female collected near 1–2 m wide creek in evergreen forest at Cham Chu Nature Reserve, Tuyen Quang
Province, Vietnam (22.2118
°
N, 105.0824
°
E, 996 m) on 27 May 2013. VNMN 2016 A.13, AMS R184836, two
adult females, and AMS R184835, adult male, collected in disturbed evergreen forest at Cham Chu Nature

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Reserve, Tuyen Quang Province, Vietnam (22.2067
°
N, 105.0783
°
E, 679 m) on 28 May 2013. AMS R184837,
adult female, and AMS R184838, AMS R184839, AMS R184841, three adult males, collected in evergreen forest
near 5–6 m wide rocky stream at Cham Chu Nature Reserve, Tuyen Quang Province, Vietnam (22.2062 N,
105.0811 E, 814 m) on 29 May 2013. AMS R184840, adult male, collected in evergreen forest at Cham Chu
Nature Reserve, Tuyen Quang Province, Vietnam (22.2054
°
N, 105.0824
°
E, 835 m) on 29 May 2013. All
specimens were collected by Jodi J. L. Rowley, Dau Quang Vinh, Hoang Duc Huy, and Hoang Anh Tuan. AMS
R184842, AMS R184843, AMS R184845, three adult females, and AMS R184844, one adult male, collected in Pe
Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2758
°
N, 103.2637
°
E, 1257 m) on 14 September
2013. VNMN 2016 A.14, adult male, and VNMN 2016 A.15, VNMN 2016 A.16, two adult females, collected in
Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2758
°
N, 103.2637
°
E, 1257 m) on 15 September
2013. VNMN 2016 A.17, VNMN 2016 A.18, VNMN 2016 A.19, three adult females, collected in Pe Xa Forest,
Sin Ho District, Lai Chau Province, Vietnam (22.2758
°
N, 103.2637
°
E, 1257 m) on 18 September 2013. VNMN
2016 A.20, one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2740
°
N,
103.2648
°
E, 1237 m) on 17 September 2013. AMS R184846, one adult female, collected in Pe Xa Forest, Sin Ho
District, Lai Chau Province, Vietnam (22.2773
°
N, 103.2626
°
E, 1260 m) on 19 September 2013. AMS R184847,
one adult female, collected in Pe Xa Forest, Sin Ho District, Lai Chau Province, Vietnam (22.2770
°
N, 103.2593
°
E, 1283 m) on 19 September 2013. AMS R184848, one adult female, collected in Pe Xa Forest, Sin Ho District,
Lai Chau Province, Vietnam (22.2687
°
N, 103.2589
°
E, 1171 m) on 21 September 2013. All specimens were
collected in evergreen forest over limestone terrain by Jodi J. L. Rowley, Le Thi Thuy Duong, Dau Quang Vinh,
Chad Minshew, and Hoang Anh Tuan.
Etymology: The specific epithet derives from the Latin petra, meaning rock, and –ops, meaning having the
appearance of, in reference to both the stone-like appearance of the frog and the limestone karst in which the
species was found in Lai Chau Province.
FIGURE 3. Male holotype (VNMN 2016 A.06) of Leptolalax petrops sp. nov. Specimen in preservative in (A) dorsal, (B)
ventral and (C) lateral view, and specimen in life in (D) dorsal and (E) ventral view. Scale bar 1mm.

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FIGURE 4. Leptolalax petrops sp. nov. in life from Tuyen Quang Province (A) Male holotype VNMN 2016 A.06 in situ, (B)
male holotype VNMN 2016 A.06, (C) paratypes AMS R184826, (D) VNMN 2016 A.09, (E) VNMN 2016 A.11 and from Lai
Chau Province (F) AMS R184845 and (G) VNMN 2016 A.16.
Diagnosis: Assigned to the genus Leptolalax on the basis of the following: small size, rounded finger tips, the
presence of an elevated thenar tubercle not continuous to the thumb, presence of macroglands on body, vomerine
teeth absent, tubercles on eyelids, anterior tip of snout with vertical white bar (Dubois 1980; Lathrop et al. 1998;
Delorme et al. 2006). The species differs from its congeners by having (1) a medium-sized body of 23.6–27.6 mm
in 21 adult males and 30.3–47.0 mm in 17 adult females, (2) immaculate white chest and belly, (3) no distinct black
markings on the head, (4) highly tuberculate skin texture, (5) toes lacking webbing and with narrow lateral fringes,
and (6) a call consisting of an average of four notes and a dominant frequency of 5.6–6.4 kHz (at 24.5–25.3 °C).
Description of holotype: Head longer than wide; snout rounded in profile and obtusely pointed in dorsal view,
projecting slightly over lower jaw; nostril approximately equidistant to tip of snout and eye; canthi rostralis
rounded, indistinct; lores sloping, slightly concave; vertical pupil; diameter of eye 84% length of snout; tympanum
distinct, round, diameter approximately half that of the eye; vomerine teeth absent; pineal ocellus absent; large oval
vocal sac openings present, located on either side of floor of mouth; tongue large, moderately broad, with wide
notch at tip; distinct, raised supratympanic ridge running from corner of eye to axillary gland. Tips of fingers

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rounded, slightly wider than width of fingers; relative finger lengths I < II < IV < III; nuptial pad absent;
subarticular tubercles absent; a large, inner palmar tubercle distinctly separated from slightly smaller, laterally
compressed outer palmar tubercle; no finger webbing or fringes. Tips of toes same width as toes; relative toe length
I < II < V < III < IV; subarticular tubercles absent, replaced by dermal ridges, distinct on second, third and fourth
toes; oval inner metatarsal tubercle pronounced, outer metatarsal tubercle absent; webbing indistinct; narrow but
distinct lateral fringes. Tibia 49% of snout-vent length; tibiotarsal articulation reaches anterior edge of eye. Skin on
dorsum mostly smooth, with indistinct, low tubercles in preservative, more obvious in life; ventral skin smooth;
dorsal ridges or dorsolateral folds absent; pectoral gland oval, approximately 1.2 mm diameter; femoral gland oval,
1.4 mm diameter, on posteroventral surface of thigh, closer to knee than to vent; supra-axillary gland raised, 1.2
mm diameter. Ventrolateral glandular line indistinct, present as indistinct, glands barely forming a line.
Colour of holotype in life: Dorsum dull medium brown with slightly darker brown markings, the most distinct
of these being a interorbital bar, a W-shaped mark on axilla, indistinct darker wash over upper tympanum, distinct
dark brown barring on upper lip, and transverse barring on dorsal surface of limbs including fingers and toes;
ventral surface of elbow and upper arm without dark bars; pale yellowish brown on elbows. Throat and ventral
surface of arms pale pink and slightly transparent, particularly at either side of throat; edges of throat pale brown
with white speckling, concentrated towards snout; ventral surface of chest and abdomen immaculate white; ventral
surface of legs pinkish grey with small whitish spots concentrated on edges. Supra-axillary gland pale yellowish
brown, pectoral gland cream, femoral glands cream. Iris gold in lower half and copper in upper half, with minute,
black reticulations.
Colour of holotype in preservative: Nearly uniform dark brown dorsal surface, with darker barring on
surface of limbs including fingers and toes. Ventral surface creamy white. Ventrolateral surfaces of lower arms and
tibiotarsus pale brown; Supra-axillary, pectoral and femoral glands creamy white.
Measurements: Holotype (mm): SVL 24.3, HDL 9.4, HDW 9.0, SNT 43.7, EYE 2.9, IOD 3.4, TMP 1.8, TEY
0.9, TIB 12.2, EN 2.1, IN 2.4, NS 1.7, ML 6.3, PL 11.3, F1 2.3, F2 3.0, F3 4.8. Weight in life 1.8 g.
TABLE 3. Measurements (mm) of adult male Leptolalax petrops sp. nov. from Tuyen Quang and Lai Chau Provinces.
Abbreviations defined in text.
Measurements Tuyen Quang Province Lai Chau Province
Range; Mean (N=19) Range; Mean (N=2)
SVL 24.3–27.6; 26.0 23.6–27.0; 25.3
HDL 9.4–10.8; 10.1 9.1–10.9; 10.0
HDW 8.9–10.1; 9.5 8.5–10.2; 9.4
SNT 2.3–4.3; 3.8 3.7–4.0; 3.9
EYE 2.9–3.7; 3.4 3.3–3.4; 3.3
IOD 2.6–3.6; 3.2 3.0–3.2; 3.1
TMP 1.7–2.3; 2.0 1.5–2.0; 1.7
TEY 0.7–1.2; 1.0 0.9–1.2; 1.1
TIB 12.0–13.5; 12.6 11.6–13.8; 12.7
EN 1.5–2.6; 2.2 2.3–2.6; 2.5
IN 2.3–2.9; 2.6 2.5–2.7; 2.6
NS 1.3–1.8; 1.6 1.4–1.7; 1.6
ML 6.3–7.8; 7.0 6.2–7.0; 6.6
PL 11.1–12.9; 11.9 10.8–12.1; 11.5
Weight (g) 1.8–2.2; 1.9 1.6–2.2; 1.9
Range; Median (N=19) Range; Median (N=2)
HDL:HDW 1.01–1.09; 1.06 1.07–1.07
HDL:SVL 0.36–0.41; 0.39 0.39–0.40
TIB:SVL 0.46–0.51; 0.48 0.49–0.51
TMP: SVL 0.06–0.09; 0.08 0.06–0.07

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TABLE 4. Measurements (mm) of adult female Leptolalax petrops sp. nov. from Tuyen Quang and Lai Chau Provinces.
Abbreviations defined in text.
Vari a t i o n: Male L. petrops sp. nov. are smaller than females (23.6–27.6 mm in 21 adult males versus 30.3–
47.0 mm in 17 adult females). The 12 females from Lai Chau Province were larger than the three from Tuyen
Quang Province (34.0–47.0 versus 30.3–33.5 mm; Table 4). Dorsal colour in life varies from pale to medium
brown, and vary in their degree of dorsal patterning. AMS R184841, AMS R184840, AMS R184831, AMS
R184827, have a nearly completely unpatterned dorsum, others have weak patterning. Individuals vary in the
number and size of dark spots along the flanks. Females are more tuberculate than males. In both sexes, skin
texture more tuberculate in life compared to in preservative. Tubercles form ridges in some individuals.
Measurements of the type series are shown in Tables 3–4.
Advertisement call: Call descriptions are based on the calls of the holotype, recorded at 24.7ºC ambient
temperature. Most calls were an average of 55 ms in duration and consisted of an average of 3.9 notes (Table 5).
Notes contained a single pulse. The dominant frequency was 6.0 kHz, and harmonics were present at
approximately 12.2 and 18.2 kHz (Fig. 5). A fundamental frequency was not evident. Calls were repeated at a rate
of 3.4 calls per second, and had an average intercall interval of 237 ms.
In six of the seven males recorded an additional call type was evident (Fig. 5Bi, iii). In the holotype, these calls
were 347 and 351 ms in duration and consisted of 24 and 23 notes of a similar frequency and amplitude to other
calls. In the calls of the seven individuals recorded, the number of notes in the most common calls varied from one
to six but four was the typical number of notes. The dominant frequency varied only slightly among individuals,
from 5.6–6.4 kHz (over 0.8 ºC difference in ambient temperature). To the human ear, the advertisement call of L.
petrops sp. nov. is a rapid, high-pitched rasping, similar to an orthopteran.
Ecology: All specimens were found at night in forest between ~234–1283 m asl. In Tuyen Quang Province,
males were observed calling from on rocks, stream banks and on vegetation within or adjacent to rocky streams in
May. During surveys in Lai Chau Province in September, no males were heard calling and most frogs encountered
were females, mostly located away from water on limestone karst.
Measurements Tuyen Quang Province Lai Chau Province
Range; Mean (N=5) Range; Mean (N=12)
SVL 30.3–33.5; 31.8 34.0–47.0; 37.9
HDL 12.0–12.8; 12.5 13.3–18.5; 14.9
HDW 11.0–12.1; 11.5 12.8–17.6; 14.0
SNT 4.3–5.0; 4.7 4.8–7.2; 5.6
EYE 3.8–4.4; 4.1 3.9–5.2; 4.5
IOD 2.9–3.6; 3.3 3.4–5.3; 4.0
TMP 2.3–2.7; 2.5 2.3–3.3; 2.7
TEY 1.0–1.5; 1.2 1.3–3.2; 1.8
TIB 14.6–15.6; 15.1 15.5–21.8; 17.5
EN 2.7–3.1; 2.9 3.0–4.3; 3.3
IN 2.8–3.4; 3.0 2.8–4.1; 3.3
NS 1.6–2.0; 1.8 1.7–2.8; 2.0
ML 7.6–8.7; 8.1 8.9–12.9; 10.2
PL 13.9–14.7; 14.3 14.9–20.9; 16.8
Weight (g) 2.5–3.1; 2.8 3.6–7.6; 4.6
Range; Median (N=5) Range; Median (N=12)
HDL:HDW 1.06–1.14; 1.09 1.02–1.09; 1.05
HDL:SVL 0.37–0.42; 0.39 0.37–0.42; 0.39
TIB:SVL 0.46–0.50; 0.47 0.43–0.50; 0.45
TMP: SVL 0.07–0.08; 0.07 0.06–0.08; 0.07

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FIGURE 5. Advertisement call of Leptolalax petrops sp. nov. (A) 30 s waveform of relative amplitude (Rel. amp.) over time
for (i) holotype VNMN 2016 A.06, (ii) paratype VNMN 2016 A.11, (iii) paratype AMS R184833, and (iv) AMS R184832. (B)
2 s waveform and corresponding spectrogram of (i) holotype VNMN 2016 A.06, (ii) paratype VNMN 2016 A.11, (iii) paratype
AMS R184833, and (iv) paratype AMS R184832. All recorded at 24.5–25.3 °C.
Conservation status: The species is currently known from Lai Chau and Tuyen Quang Provinces, Vietnam.
The true extent of occurrence is unknown but the species probably extends further into adjoining areas of
northwestern Vietnam and southern China. Suitable forested regions may include parts of Hoang Lien Son Nature
Reserve, Muong Nhe Nature Reserve and Nam Don Nature Reserve in Vietnam. We recommend that the species is
listed as Vulnerable on the Red List of Threatened Species as its predicted Extent of Occurrence is 19,874km
2
, it
likely occurs in only two threat-defined locations, and there is an observed continuing decline in the quality of its
habitat, satisfying Red List Criteria B1ab(iii).

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TABLE 5. Measurements of advertisement call parameters for Leptolalax petrops sp. nov. from Tuyen Quang Province.
Parameter values are given as means (and ranges). *holotype.
Comparisons: Leptolalax petrops sp. nov. differs morphologically and/or bioacoustically to all known
Leptolalax species. From species in the subgenus Lalos (Dubois et al. 2010), distributed south of the Isthmus of
Kra, Leptolalax petrops sp. nov. differs by lacking ventrolateral glands (present in the subgenus Lalos; Delorme et
al. 2006) and by molecular divergence (Table 2, Fig. 2).
From the 35 Leptolalax species considered to be in subgenus Leptolalax (Delorme et al. 2006) or in the
Leptolalax applebyi group (subgeneric placement remains unresolved; Rowley et al. 2016), Leptolalax petrops sp.
nov. differs by a combination of morphological, molecular and/or bioacoustic data. From the 23 species with
molecular data available, the new species differs in a high divergence in the 16S gene fragment examined (Table 2,
Fig. 2), and by a combination of morphological and bioacoustic characters. From L. aureus, the new species differs
by having highly tuberculate dorsal skin (versus finely tuberculate), having black lateral spots (versus no lateral
spots), and a call consisting of 1–6 notes (average of four) (versus a call consisting of 2–16 notes). From L.
applebyi, the new species differs by having a body size of 23.6–27.6 mm in 21 adult males and 30.3–47.0 mm in 17
adult females (versus 19.6–22.3 mm in males and 21.7–25.9 mm in females), having highly tuberculate dorsal skin
(versus mostly smooth), an immaculate white chest and belly (versus reddish brown with white speckling), lacking
distinct black markings on the head (versus black markings present), and a call with a dominant frequency of 5.6–
6.4 kHz at 24.5–25.3 °C (versus 4.0–4.3 kHz at 21.5 °C). From L. ardens, the new species differs by having highly
tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus reddish brown with
white speckling), lacking distinct black markings on the head (versus black markings present), and a call with a
dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C (versus 3.1–4.3 kHz at 21.5 °C). From L. bidoupensis, the new
species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and
belly (versus reddish brown with white speckling), lacking distinct black markings on the head (versus black
markings present), and a call consisting of an average of four notes (versus 6–9 notes) and a dominant frequency of
5.6–6.4 kHz at 24.5–25.3 °C (versus 1.9–3.8 kHz at 19–21 °C). From L. bourreti, the new species differs by having
a body size of 23.6–27.6 mm in adult males (versus 36.2 mm), highly tuberculate dorsal skin (versus mostly
smooth), and lacking distinct black markings on the head (versus black markings present). From L. eos, the new
species differs by having a male body size of 23.6–27.6 mm (versus 33.1–34.7 mm), and having highly tuberculate
AMS
R184826
VNMN 2016
A.06*
VNMN
2016 A.09
VNMN
2016 A.11
AMS
R184831
AMS
R184832
AMS
R184833
Primary
call
Call duration
(ms)
42.6 (10–
57)
55.0
(44–57)
52.1
(5–56)
55.5 (51–
60)
53.2 (50–
55)
54.8 (52–
58)
59.9 (57–
62)
Intercall interval 416 (337–
660)
237.4 (174–
273)
296.3 (175–
637)
408.3 (317–
568)
325.3
(286–530)
261.2
(232–375)
537.2
(299–805)
Notes/call 4.1 (1–6) 3.9 (3–4) 4.1 (4–5) 4 4 4 4
Call repetition
rate
2.2 3.4 2.8 2.1 2.6 3.1 1.7
Fundamental
frequency
0.2 0.2 0.2 0.2 0.2 0.2 0.2
Dominant
frequency
5.7 (5.6–
5.8)
6 6.0 (5.8–
6.0)
5.8 (5.8–
6.0)
6.3 (6.2–
6.4)
6.1 (6.0–
6.4)
5.8
Harmonic 1 11.4 12.2 12 11.6 12.6 12.1 11.6
Harmonic 2 17.1 18.2 17.6 18.4 18.6
Harmonic 3 23.6 24.6
Secondar
y call
Duration none 347, 351 379 228, 266 215 378, 354 217, 196
Notes/call 24, 23 28 16, 18 18, 21, 21 14, 13
Note repetition rate 66, 63 71 66, 64 79, 53, 56 60, 61
Dominant frequency (kHz) 6.6 5.8 5.3, 5.8 6.4 5.8, 5.8 5.8, 5.6
Temperature (º C) 24.5 24.7 25.1 24.7 25.3 25.2 24.8

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dorsal skin (versus shagreened). From L. firthi, the new species differs by having black lateral spots (versus no
lateral spots), feet with narrow but distinct lateral fringes (versus feet with wide lateral fringes in males), and a call
repetition rate of 1.7–3.1 calls/s at 24.5–25.3 °C (versus <1 call/s at 21.5 °C). From L. isos, the new species differs
by having black lateral spots (versus no lateral spots), feet with narrow but distinct lateral fringes in both sexes
(versus feet with wide lateral fringes in males), and a call consisting of an average of four notes (versus three
notes). From L. kalonensis, the new species differs by having highly tuberculate dorsal skin (versus mostly
smooth), an immaculate white chest and belly (versus dark brownish with white speckling), and a call with a
dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C (versus 2.8 kHz at 26.4 °C). From L. laui, the new species
differs by having highly tuberculate dorsal skin (versus shagreened), and feet with narrow but distinct lateral
fringes in both sexes (versus feet with wide lateral fringes). From L. liui, the new species differs by having highly
tuberculate dorsal skin (versus mostly smooth), lacking distinct black markings on the head (versus black markings
present), and feet with narrow but distinct lateral fringes in both sexes (versus feet with wide lateral fringes). From
L. maculosus, the new species differs by having highly tuberculate dorsal skin (versus mostly smooth), an
immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on
the head (versus black markings present), and a call with a dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C
(versus 2.7–2.8 kHz at 23.3–24.1 °C). From L. melicus, the new species differs by having a male body size of 23.6–
27.6 mm (versus 19.5–22.7 mm in males), highly tuberculate dorsal skin (versus mostly smooth), an immaculate
white chest and belly (versus reddish brown with white speckling), lacking distinct black markings on the head
(versus black markings present), and a call with a dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C (versus 2.9–
3.8 kHz at 26.1–26.2 °C). From L. minimus, the new species differs by having highly tuberculate dorsal skin
(versus mostly smooth). From L. nyx, the new species differs by lacking dark, regularly set rounded spots on the
dorsum (versus dark, regularly set rounded spots present), and highly tuberculate dorsal skin (versus mostly
smooth). From L. oshanensis, the new species differs by having highly tuberculate dorsal skin (versus smooth with
fine glandular ridges), no distinct black markings on the head (versus black markings present), and a call with an
average of four short repeated notes (versus three) and a dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C
(versus 4.4–4.6 kHz at 14 °C). From L. pallidus, the new species differs by having an immaculate white chest and
belly (versus brown with white speckling), and a call with a dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C
(versus 2.4–2.7 kHz at 14.0–21.4 °C). From L. pyrrhops, the new species differs by having a male body size of
23.6–27.6 mm (versus 30.8–34.3 mm), highly tuberculate dorsal skin (versus finely shagreened), an immaculate
white chest and belly (versus brownish with white speckling), no distinct black markings on the head (versus black
markings present), and a call with an average of four notes (versus 5–6 notes) and a dominant frequency of 5.6–6.4
kHz at 24.5–25.3 °C (versus 1.9–2.2 kHz at 21.5 °C). From L. pluvialis, the new species differs by having a male
body size of 23.6–27.6 mm (versus 21.3–22.3 mm), a highly tuberculate dorsal skin (versus mostly smooth), an
immaculate white chest and belly (versus dirty white with dark brown marbling), and no distinct black markings on
the head (versus black markings present). From L. tadungensis, the new species differs by having highly
tuberculate dorsal skin (versus smooth), an immaculate white chest and belly (versus brown with white speckling),
lacking distinct black markings on the head (versus black markings present), and a call with a dominant frequency
of 5.6–6.4 kHz at 24.5–25.3 °C (versus 2.6–3.1 kHz at 12.9–22.3 °C). From L. tengchongensis, the new species
differs by having an immaculate white chest and belly (versus with dark brown blotches on chest and belly), and
lacking distinct black markings on the head (versus black markings present). From L. ventripunctatus, the new
species differs by having highly tuberculate dorsal skin (versus mostly smooth with longitudinal ridges), an
immaculate white chest and belly (versus with dark spots), and lacking distinct black markings on the head (versus
black markings present). From L. zhangyangpingi, the new species differs by having a male body size of 23.6–27.6
mm (versus 45.8–52.5 mm) and highly tuberculate dorsal skin (versus mostly smooth with distinct tubercles and
longitudinal ridges).
From the remaining 12 Leptolalax species in the subgenus Leptolalax (Delorme et al. 2006), no molecular data
are available and their genetic differentiation to Leptolalax petrops sp. nov. could therefore not be assessed,
however they differ by a combination of morphological and bioacoustics characters. From L. alpinus, the new
species differs by having highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and
belly (versus dark spotting), lacking distinct black markings on the head (versus black markings present), and a call
consisting of an average of four notes (versus an average of 9.5 notes). From L. botsfordi, the new species differs
by having a male body size of 23.6–27.6 mm (versus 29.1–32.6 mm), having highly tuberculate dorsal skin (versus

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shagreened), an immaculate white chest and belly (versus reddish brown with white speckling), lacking distinct
black markings on the head (versus black markings present), and a call consisting of an average of four notes
(versus 1–3 notes) and a dominant frequency of 5.6–6.4 kHz at 24.5–25.3 °C (versus 2.6–3.4 kHz at 14 °C). From
L. croceus, the new species differs by having an immaculate white chest and belly (versus bright orange belly), a
visible tympanum (versus indistinct tympanum), and a call consisting of an average of four short repeated notes
(versus 4–6 highly variable notes). From L. fuliginosus, the new species differs by having a body size of 23.6–27.6
mm in adult males (versus 28.2–30.0 mm), highly tuberculate dorsal skin (versus mostly smooth), an immaculate
white chest and belly (versus dusty), and lacking distinct black markings on the head (versus black markings
present). From L. khasiorum, the new species differs by having, an immaculate white chest and belly (versus venter
with darker pigments on sides), and an iris that is gold in lower half and copper in upper half (versus iris with bright
orange upper). From L. lateralis, the new species differs by having highly tuberculate dorsal skin (versus finely
tuberculate), a head longer than wide (versus HW>HL), and nostril closer to the snout than eye (versus closer to
eye). From L. nahangensis, the new species differs by having a body size of 23.6–27.6 mm in adult males (versus
male 40.8 mm), highly tuberculate dorsal skin (versus mostly smooth), an immaculate white chest and belly (versus
light specking on throat and chest), and lacking distinct black markings on the head (versus black markings
present). From L. nokrekensis, the new species differs by having a highly tuberculate dorsal skin without
longitudinal folds (versus dorsum with longitudinal folds), lacking distinct black markings on the head (versus
black markings present) and an iris that is gold in lower half and copper in upper half (versus iris with bright red
upper). From L. pelodytoides, the new species differs by having a highly tuberculate dorsal skin (versus mostly
smooth with small, smooth warts) and feet with indistinct webbing (versus feet one-third webbed). From L. sungi,
the new species differs by having a male body size of 23.6–27.6 mm (versus 48.3–52.7 mm), highly tuberculate
dorsal skin (versus uniformly finely granular), medium brown dorsum (versus pale creamy brown), and lacking
distinct black markings on the head (versus black markings present). From L. tamdil, the new species differs by
having a body size of 23.6–27.6 mm in adult males (versus male 32.3 mm), highly tuberculate dorsal skin (versus
weakly tuberculate) and an iris that is gold in lower half and copper in upper half (versus top third bright orange,
lower greyish-cream). From L. tuberosus, the new species differs by having an immaculate white chest and belly
(versus with black streaks), having a distinct tympanum (versus indistinct), a call with an average of four notes
(versus a single note).
Discussion
The incorporation of molecular and bioacoustic data has been instrumental in facilitating species discovery in this
morphologically conserved genus. In the last three years alone, 11 new Leptolalax species have been described
(Matsui et al. 2014a, Matsui et al. 2014b, Sung et al. 2014, Rowely et al. 2015, Poyarkov et al. 2015, Rowley et al.
2016, Yang et al. 2016). The description of Leptolalax petrops sp. nov. brings the total number of known species in
the genus to 51.
We consider that specimens from both Tuyen Quang and Lai Chau provinces belong to the new species. The
genetic divergence between the sites (2.7–3.1% divergence at the 16S gene fragment analysed), located over 180
km apart, is less than that between any known Leptolalax species for which comparable sequences are available.
Sampling from between these localities, and the incorporation of bioacoustics data from Lai Chau Province is
needed to confirm their placement.
Most species of Leptolalax appear to be dependent on forest, making ongoing deforestation in the region
(Meyfroidt & Lambin 2008) a significant threat to the genus, as it is for other frog species in the region (Rowley et
al. 2010d). Indeed, we observed forest loss at both known localities, but particularly at the site in Lai Chau
Province (Fig. 6B). Further research is needed to understand this species’ true range and threats, and there is an
urgent need to assess the conservation status of species as soon as possible after description. As such, we have
provided information to assist its assessment on the IUCN Red List of Threatened Species and we recommend that,
as much as possible, such information should be provided in new species descriptions to facilitate the prioritization
of conservation resources.

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FIGURE 6. Habitat of Leptolalax petrops sp. nov. in (A–B) Cham Chu Nature Reserve, Tuyen Quang Province, and (C–D) Pe
Xa Forest, Lai Chau Province.
Acknowledgements
Research was supported by grants from the ADM Capital Foundation and National Geographic Committee for
Research and Exploration, and by a grant from the President of the Vietnam Academy of Science and Technology
(VAST.CTG.03/16-17) to T.T. Nguyen. Prof. Nguyen Trung Minh and Prof. Luu Dam Cu (Vietnam National
Museum of Nature, Hanoi) supported the collaboration and fieldwork. Staff at Cham Chu Proposed Nature Reserve
assisted during fieldwork and issued permissions (Permit numbers 489/TCLN-BTTN from the Vietnamese
Ministry of Agriculture and Rural Development and 1110/UBND-NLN from Tuyen Quang province). Chad
Minshew and Hoang Anh Tuan assisted in the field. For all this assistance we are most grateful.

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APPENDIX I. Comparative material examined.
Leptolalax aereus: Laos, Savannakhet Province, Vilabouli District (NCSM 76038–76057, 76061–76071; SAMA R64234–
64242; type specimens).
Leptolalax applebyi: Vietnam, Quang Nam Province, Song Thanh Nature Reserve (AMS R171703–171707; type specimens).
Vietnam, Kon Tum Province, Ngoc Linh Nature Reserve (AMS R173635, 177637, UNS 00462/AMS R173776, AMS
R173777, AMS R173778, UNS 00465/AMS R173780, AMS R176528, UNS 00464/AMS R176529, UNS 00465/AMS
R176530, AMS R176531–176533).
Leptolalax ardens, Vietnam, Gia Lai Province (VNMN 04707/AMS R176454, VNMN 04708/AMS R176455, VNMN 04709/
AMS R176456, VNMN 04710/AMS R176457, VNMN 04711/AMS R176458, VNMN 04712/AMS R176459; type
specimens).
Leptolalax bidoupensis: Vietnam, Lam Dong Province, Bidoup-Nui Ba National Park (AMS R173133, UNS 00101/AMS
R173135, UNS 00102/AMS R173137, AMS R173134, AMS R173136, NCSM 77320, NCSM 77321, NCSM 77322; type
specimens). Vietnam, Lam Dong Province, Bidoup-Nui Ba National Park (UNS00522, ZFMK 96595, UNS00523, ZFMK
96596, UNS00524, ZFMK 96597, UNS00525).
Leptolalax botsfordi: Vietnam, Lao Cai Province, Hoang Lien National Park (VNMN 03682, AMS R176534–176541; type
specimens).
Leptolalax croceus: Vietnam, Kon Tum Province, Ngoc Linh Nature Reserve (AMS R173738–173748, UNS 00108/ AMS
R173750, AMS R173751, UNS 00109/AMS R173752, UNS 00110/AMS R173775, UNS 00111/AMS R173779; type
specimens).
Leptolalax dringi: Malaysia, Sarawak (AMNH A90487).
Leptolalax eos: Laos, Phongsaly Province (FMNH 258052–258064, 258066). Laos, Luang Namtha Province (FMNH 271281–
271285).
Leptolalax firthi: Vietnam, Kon Tum Province, Ngoc Linh Nature Reserve (AMS R176524, AMS R176525, UNS 00460/AMS
R176526, UNS 00461/AMS R176527, AMS R176513, UNS 00456/AMS R176514, AMS R176515, NCSM 78995, AMS
R176503–176507, UNS 00453/AMS R176509, UNS 00454/AMS R176510, UNS 00455/AMS R176511, AMS R176500,
UNS 00452/AMS R176501, AMS R176502, AMS R176508, AMS R1765012, AMS R176517, NCSM 78996–78997,
AMS R176518, UNS 00457/AMS R176516, AMS R176520, AMS R176522, UNS 00458/AMS R176519, AMS
R176521, UNS 00459/AMS R176523, AMS R173736, AMS R173774). Vietnam, Quang Nam Province, Song Thanh

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Proposed Nature Reserve (AMS R171714, AMS R171722)
Leptolalax gracilis: Malaysia, Sarawak (AMNH A90488).
Leptolalax isos: Vietnam, Gia Lai Province, Kon Ka Kinh National Park (VNMN A 2015.4/AMS R 176480, VNMN A.2015.5/
AMS R 176491, A.2015.6/AMS R 176492, AMS R 176468–176479, 176481–176485-90, 176493, 176494, NCSM 79651,
79652; type specimens). Cambodia Ratanakiri Province, Virachey National Park (MVZ 258072, 258073, 258184–6,
258188, 258190–258196; type specimens).
Leptolalax kalonensis: Vietnam, Binh Thuan Province (IEBR A.2014.14–A.2014.22; type specimens).
Leptolalax liui: China, Fukien Province, Ch’ungsan, Hsein (AMNH A30127, 30129, 30130, 30135; FMNH 22415–22418;
topotypes). China, Fukien Province, Yenping (AMNH A28417).
Leptolalax maculosus: Vietnam, Ninh Thuan Province (ZFMK 96600, UNS00513, UNS00514; type specimens).
Leptolalax melicus: Cambodia, Ratanakiri Province, Virachey National Park (MVZ 258074–258077, MVZ 258197– 258199;
type specimens).
Leptolalax nyx: Vietnam, Ha Giang Province (AMNH A163798, 163799, 163802–163804, 163811–163813, 163817–163819,
163822–163824, 163828–163830; topotypes).
Leptolalax oshanensis: China, Sichuan Province, Hongya Xian (FMNH 232907–232921; ~50 km from type locality).
Leptolalax pallidus: Vietnam, Lam Dong Province (UNS00510–00512, ZFMK 96598, ZFMK 96599; type specimens).
Leptolalax sungi: Vietnam, Lao Cai Province (AMNH 168676).
Leptolalax tadungensis: Vietnam, Dak Nong Province (UNS00515–00517, UNS 00526, UNS 00527, ZFMK 96601–96603).
Leptolalax tuberosus: Vietnam, Quang Nam Province, Song Thanh Nature Reserve (AMS R171715–171721). Vietnam, Quang
Nam Province (AMNH A163665).
Leptolalax ventripunctatus: Laos, Phongsaly Province (FMNH 258077). Laos, Luang Namtha Province (FMNH 271317–21,
271329, 271331).