440 Accepted by J. Rowley: 26 Nov. 2013; published: 9 Jan. 2014
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3753 (5): 440–452
A new Leptolalax from the mountains of Sabah, Borneo
(Amphibia, Anura, Megophryidae)
MASAFUMI MATSUI1,3, KANTO NISHIKAWA1 & PAUL YAMBUN2
1Graduate School of Human and Environmental Studies, Kyoto University, Kyoto 606-8501, Japan
2Research and Education Division, Sabah Parks, P.O. Box 10626, Kota Kinabalu 88806, Sabah, Malaysia
3Corresponding author. E-mail: email@example.com
A new species of Leptolalax is described from Kinabalu National Park in western Sabah, Malaysian Borneo. The new
species had been assigned to L. dringi, L. gracilis, or L. fritinniens in the past. It differs from all congeners, including these
species, by a unique combination of morphological characters, including small body size, rounded snout, narrower inter-
orbital than upper eyelid, basal toe webbing, smooth skin with tiny tubercles on dorsum and dorsal side of head, small
pectoral glands, absence of supraaxillary glands and ventrolateral glandular ridges, spotted venter, advertisement call con-
sisting of long series of 1–149 notes, each composed of three or four pulses, and dominant frequency at 6.90–7.35 kHz,
without prominent frequency modulation.
Key words: Amphibia, Leptolalax sabahmontanus sp. n., advertisement call, Sabah, mtDNA phylogeny
The megophryid genus Leptolalax Dubois occurs widely in the Southeast Asia and seven species are currently
recorded from Borneo, and several additional cryptic species remain to be described (Dehling & Matsui 2013).
Acoustic characteristics are effective in identifying species of Leptolalax (e.g. Malkmus & Riede 1993; Matsui
1997, 2006; Matsui et al. 2009; Rowley & Cao 2009; Rowley et al. 2012), and additionally, DNA barcoding (e.g.,
Vences et al. 2005) is now becoming popular, enabling taxonomic works on this genus to be more accurate and
easier (Ohler et al. 2011).
Leptolalax dringi Dubois was originally described from Gunung (= Mt.) Mulu, Sarawak, western Borneo of
East Malaysia (Matsui & Dehling 2012), and has also been reported from Mt. Kinabalu of Sabah, northern Borneo
(Malkmus et al. 2002), although their species identity is doubted (Matsui & Dehling 2012). Our field survey and
examination of museum specimens revealed the presence on the Crocker range, south of Mt. Kinabalu in Sabah, of
specimens resembling the Kinabalu specimens. Subsequent morphological and genetic analyses revealed that
populations from Sabah (Kinabalu and Crocker) substantially differ from L. dringi from Sarawak in body size and
some body proportions, calls, and mtDNA gene sequences. We therefore describe the population from Sabah as a
Material and methods
For adult specimens stored in 70% ethanol, we took the following 19 body measurements to the nearest 0.1 mm
under a binocular dissecting microscope, following Matsui (1984): (1) snout-vent length (SVL); (2) head length
(HL); (3) head width (HW); (4) internarial distance (IND); (5) interorbital distance (IOD); (6) upper eyelid width
(UEW); (7) snout length (SL); (8) eye length (EL); (9) nostril-eyelid length (N–EL); (10) tympanum diameter
(TD); (11) tympanum-eye length (T-EL); (12) lower arm and hand length (LAL); (13) forelimb length (FLL); (14)
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NEW MONTANE LEPTOLALAX FROM BORNEO
inner palmar tubercle length (IPTL); (15) tibia length (TL); (16) foot length (FL); (17) hindlimb length (HLL); (18)
inner metatarsal tubercle length (IMTL); and (19) first toe length (1TOEL). We followed the system of description
of toe-webbing states used by Savage & Heyer (1967). Specimens are deposited at the Institute for Tropical
Biology and Conservation, University Malaysia Sabah (BORNEENSIS), Sabah Parks (SP) and the Graduate
School of Human and Environmental Studies, Kyoto University (KUHE).
For the acoustic data, we recorded frog calls in the field using a Sony TC-D5M stereo cassette recorder with a
Sony ECM-23F microphone (the recordings of which were digitized at 16 bits and 44.1 kHz) and analyzed them
with SoundEdit Pro (MacroMind-Paracomp, Inc.) and Raven Lite 1.0 for Mac OS X (http://www.birds.cornell.edu/
raven) on a Macintosh computer. Temporal data were obtained from the oscillogram and frequency information
was obtained from the audiospectrograms using Fast Fourier Transformation (1024 point Hanning window).
We obtained DNA sequence data from tissue samples preserved in 99% ethanol. We reconstructed
phylogenetic trees from approximately 1900 base pairs (bp) of the partial sequences of mitochondrial 12S and 16S
rRNA genes to clarify the genetic structure of our own samples of three specimens of Leptolalax sp. from Sabah
(BORNEENSIS 12632 from Liwagu, Kinabalu; BORNEENSIS 22784 and 12454 from Mahua, Crocker), three
topotypic samples of L. dringi from Camp 4 of Gunung Mulu (KUHE 55610, 55612, 55613), each one sample of L.
fritinniens Dehling & Matsui (KUHE 55371 from Headquarters, Gunung Mulu), L. pictus Malkmus (KUHE 39298
from Poring, Kinabalu), L. maurus Inger, Lakim, Biun & Yambun (SP 21450 from Kinabalu), L. arayai Matsui
(BORNEENSIS 22931 from Liwagu, Kinabalu), and L. gracilis (Günther) (KUHE 55624 from Camp 1, Gunung
Mulu), and outgroup species L. heteropus (Boulenger) (KUHE 15490 from Larut, Peninsular Malaysia) and
Leptobrachium hasselti Tschudi (KUHE 42820 from Java: AB530424). Methods for DNA extraction and,
amplification and sequencing of the mtDNA fragments are the same as those reported by Matsui et al. (2010) and
Hamidy et al. (2011). Briefly, we amplified DNA fragments by PCR with the primers detailed in Hamidy et al.
(2011) and sequenced amplified fragments using the amplification primers and 10 additional sequencing primers.
The PCR cycling, precipitation, and sequencing procedures were identical to those described by Hamidy et al.
(2011). The resultant sequences were deposited in GenBank (AB847550–847561). The alignment matrix with
1935 mtDNA nucleotide sites (529 sites for 12S rRNA; 1406 for 16S rRNA) was subjected to estimate
phylogenetic relationships using maximum likelihood (ML) and Bayesian inference (BI). Pairwise comparisons of
uncorrected sequence divergences (p-distance) were also calculated for 16S rRNA. Details for all these procedures
are given in Hamidy et al. (2011).
As a result of molecular phylogenetic analyses, we obtained 1907 bp of concatenated fragments of mtDNA genes
for all samples, including out-groups (Figure 1). Of 1907 nucleotide sites, 482 were variable, and 253 were
parsimoniously informative. The best substitution model was J2 with gamma shape parameter (G) of 0.346 for ML
and GTR+G of 0.370 for BI. The likelihood values (-lnLs) of the ML and BI trees were 10010.084 and 10027.834,
Phylogenetic analyses employing two different optimality criteria yielded identical relationships. As shown in
the ML tree in Figure 1, three specimens of Leptolalax sp. from Sabah examined here proved to form a well-
supported clade (MLBS = 90, Bayesian posterior probability [BPP] = 1.00), which was sister to the clade of L.
dringi with high support values (MLBS = 99, BP P = 1.00) (Fig. 1). The genetic distance (uncorrected p-distance of
16S rRNA) between these two clades was substantially large, with the mean of 5.5% (range = 5.1–5.8%: Table 1).
This value was only slightly smaller than the distance between distinctly different species L. dringi and L. pictus
(5.7–6.3%, mean = 6.1%).
Furthermore, in congruence with genetic separation, the specimens of Leptolalax sp. from Sabah are also
clearly separated morphologically from all the other congeners, including its sister species L. dringi. We thus
describe Leptolalax sp. from Sabah as follows:
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TABLE 1 . Uncorrected p-distances (in %) for fragment of 16S rRNA among Leptolalax taxa compared.
FIGURE 1. Maximum likelihood (ML) tree of a 1907 bp sequence of mitochondrial 12S and 16S rRNA for samples of
Leptolalax (for sample details, see text). Numbers above or below branches represent bootstrap supports for ML inference and
Bayesian posterior probability (ML-BS/BPP).
Leptolalax sabahmontanus sp. nov.
(Figures 2, 3, 4)
Holotype. BORNEENSIS 22684, an adult male from Silau Silau trail along Sungai Silau-Silau (06o00'N, 116o32'E,
alt. 1450 m a.s.l.), Kinabalu National Park (KNP), Ranau District, Sabah, Malaysia, collected on 31 July 2005 by
Paratypes. BORNEENSIS 22683, 22685, 22687–22691, 22701, KUHE 48014–48016 (former
BORNEENSIS 22681, 22682, 22686), 11 males data, same as the holotype. BORNEENSIS 22757, 22760, KUHE
48017 (former BORNEENSIS 22756), three males from the type locality, collected on 2 August 2005 by M.
1Leptolalax sp. BORNEENSIS 22784 (AB847552)
2Leptolalax sp. BORNEENSIS 12632 (AB847551) 0.0
3Leptolalax sp. BORNEENSIS 12454 (AB847550) 0.1 0.1
4L. dringi KUHE 55610 (AB847553) 5.1 5.1 5.2
5L. dringi KUHE 55612 (AB847554) 5.7 5.7 5.8 0.6
6L. dringi KUHE 55613 (AB847555) 5.7 5.7 5.8 0.6 0.0
7L. pictus KUHE 39298 (AB847556) 6.3 6.3 6.3 5.7 6.3 6.3
8L. furitinniens KUHE 55371 (AB847557) 7.2 7.2 7.3 7.4 7.9 7.9 6.7
9L. arayai BORNEENSIS 22931 (AB847558) 11.9 11.9 11.9 11.9 12.4 12.4 10.8 11.7
10 L. maurus SP 21450 (AB847559) 12.1 12.1 12.1 12.5 13.1 13.1 11.8 12.4 10.0
11 L. gracilis KUHE 55624 (AB847560) 15.1 15.1 15.1 14.8 15.4 15.4 14.2 15.6 15.7 15.4
12 L. heteropus KUHE 15490 (AB847561) 19.8 19.8 19.8 19.9 20.5 20.5 19.3 19.2 21.3 21.0 21.4
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NEW MONTANE LEPTOLALAX FROM BORNEO
Matsui, P. Yambun, K. Nishikawa, and T. Shimada. SP 26736, 26737, two males from the type locality, collected
on 14 August 2008 by Majin Silim. BORNEENSIS 22748, 22749, two males from Pandanus trail (06o00'N,
116o32'E, 1500 m a.s.l.), KNP, collected on 1 August 2005 by M. Matsui, P. Yambun, K. Nishikawa, and T.
Shimada. KUHE 48018 (former BORNEENSIS 22784), BORNEENSIS 22785, 22786, three males from junction
of Sg. Liwagu and Sg. Silau-Silau (06o00’ N, 116o33’ E, 1400 m a.s.l.), KNP, collected on 5 August 2005 by M.
Matsui, P. Yambun, K. Nishikawa, and T. Shimada. SP 26740, an adult female from Bundu Tuhan view trail, KNP,
collected on 14 August 2008 by S. David. BORNEENSIS 22980, 23029, 23034, three males from Sayap (06o11’ N,
116o34’ E, 950m a.s.l.), KNP, Kota Belud District, collected on 12 August 2005 by M. Matsui, Y. B. Paul, K.
Nishikawa, and T. Shimada.
Referred specimens. BORNEENSIS 8711, 8726, 8729, three males from Trail 4 (05o29’ N, 116 o01’ E, 754 m
a.s.l.) of UMS 2002 expedition to the Crocker Range, Ulu Kimanis, Crocker Range National Park (CRNP),
Tambunan District, collected on 9 September 2002 by M. Matsui. BORNEENSIS 8179, one male from Trail 5
(05o29’ N, 116o01’ E, 820 m a.s.l.) of Ulu Kimanis, CRNP, collected on 1 September 2002 by M. Matsui.
BORNEENSIS 8673, one male collected on 7 September 2002 by M. Matsui, and KUHE 48020 (former
BORNEENSIS 23470), one male collected on 19 August 2005 by M. Matsui, from Trail 11 (05o27’ N, 116o03’ E,
1427 m a.s.l.) of Ulu Kimanis, CRNP. KUHE 48019 (former BORNEENSIS 8855), one male collected on 12
December 2003, BORNEENSIS 12454, one male collected on 5 August 2003 by M. Matsui, BORNEENSIS
12632–12634, three males collected on 9 August 2003 by M. Matsui, K. Nishikawa, and T. Shimada, and
BORNEENSIS 23482, one male collected on 20 August 2005 by M. Matsui, all from Mahua, CRNP (05o48’ N,
116o24’ E, 1063 m a.s.l.).
FIGURE 2. Dorsolateral view of a male paratype of Leptolalax sabahmontanus sp. nov. (KUHE 48014) from the type locality,
Silau Silau trail along Sg. Silau-Silau, Kinabalu National Park.
Etymology. The specific epithet is from Sabah and a Latin adjective montanus denoting mountain, alluding to
the fact that the species inhabits on mountains of Sabah, northern Borneo.
Diagnosis. The new species is assigned to Leptolalax on the basis of the following: lack of vomerine teeth;
fingers rounded in tips and lacking webbing; inner palmar tubercle elevated, not extending to the first finger;
subarticular tubercles indistinct, replaced by low callous tissue; lack of nuptial pad; toe webbing basal; lack of
outer metatarsal tubercle; snout tip with white vertical bar (Dehling & Matsui 2013). A medium-sized form of the
genus, adult males about 25–28 mm in SVL; snout rounded in both ventral view and lateral view; interorbital space
narrower than upper eyelid; male vocal sac not bipartite; toe webbing basal; dorsal skin of trunk and head smooth
with tiny tubercles; pectoral gland small and indistinct; supraaxillary gland and ventrolateral glandular ridges
absent; venter spotted; advertisement call consisting of long series of 1–149 notes, each composed of one to two
indistinct pulse groups, dominant frequency at 6.75–7.35 kHz, without prominent frequency modulation. Most
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similar to L. dringi, but differing in smaller body size, relatively smaller eye-tympanum distance and inner
metatarsal tubercle, and larger interorbital distance, upper eyelid width, snout length, eye length, and forearm and
hand length compared to L. dringi.
Description of holotype (measurement in mm). SVL 26.3; head and body moderately stocky (Figures 3);
head longer (10.7) than broad (9.7) and wider than trunk; snout rounded in both ventral view and lateral view,
slightly protruding beyond lower jaw; eye large (4.1) subequal to snout length (4.0); pupil vertical; canthus rostralis
distinct, slightly concave in dorsal view between eye and nostril; convex between nostril and tip of snout; loreal
region oblique, concave; nostril lateral, below canthus, distinctly closer to eye (2.1) than to tip of snout; internarial
space (2.5) narrower than interorbital distance (3.0), latter slightly narrower than upper eyelid (3.2); pineal spot
invisible; tympanum distinct, rounded, diameter (2.4) three-fifths that of eye, and separated from eye by about one-
third tympanic diameter (0.9); symphysial knob on anteriormost part of mandible; vomerine ridge and teeth absent;
tongue large, broad, notched distally, without papillae; a median, subgular vocal sac in the mid-ventral line; vocal
openings small, just posterior to rictus, close to eustachian tube openings.
FIGURE 3. Dorsal (A) and ventral (B) views of male holotype of Leptolalax sabahmontanus sp. nov. (BORNEENSIS 22684).
Scale bar = 10 mm.
Forelimb slender, moderately long (19.7); hand and forearm long (15.0); fingers long and slender, without
webbing or lateral fringe of skin (Figure 4A); relative length of fingers II<I< IV <III; tips rounded and thickened;
nuptial pad absent; inner metacarpal tubercle large, subquadratic, and prominent in thenar and metacarpal region of
fingers I and II, divided subequally in proximal and distal part by shallow superficial depression, separated by a
distinct groove from much smaller, outer metacarpal tubercle in metacarpal region of fingers III and IV; subarticular
tubercles indistinct, replaced by long, low callous tissue.
Hindlimb relatively long (43.7), more than two times length of forelimb; tibia (14.0) longer than foot (12.2); heels
overlapping when limbs are held at right angles to body; tibiotarsal articulation of adpressed limb reaching level
between nostril and snout; relative length of toes I<II<V<III<IV; toe tips rounded and thickened, smaller than finger
tips; toe webbing basal, webbing formula I 13/4–2+ II 2–3+ III 3–4 IV 4+–23/4 V (Figure 4B); fringes of skin on lateral
sides of toes very narrow, hardly discernible; inner metatarsal tubercle low, oval, large, length (1.4) slightly more than
half length of first toe (2.0); no outer metatarsal tubercle; subarticular tubercles obscure, but elongate, replaced by low
callous tissue from base to distal half of penultimate phalanx.
Skin on dorsum and dorsal side of head scattered with tiny tubercles, weakly wrinkled on dorsal surfaces of limbs
(Figures 3A); lateral surface of trunk with an irregular row of several larger, low tubercles; weak wrinkles on dorsal
surfaces of limbs forming indistinct, reticulated, predominantly longitudinal, low ridges; supratympanic fold low but
distinct, slightly angular, running from posterior margin of eye to level of insertion of forelimb; several enlarged
tubercles at posterior end of rictus; chest and abdomen smooth, skin of gular region not modified; pectoral gland very
small and low, hardly discernible, at insertion of forelimb; supraaxillary gland hardly discernible, and femoral gland
and ventrolateral glandular ridges absent.
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NEW MONTANE LEPTOLALAX FROM BORNEO
Colour. In life, olive brown dorsally on head and body, with large, dark brown, irregularly shaped markings,
some with trace of light outline; medial dark brown markings comprising small reverse-triangle pattern on snout,
much larger, similar-shaped one between eyes and continuous W-shaped pattern on shoulder region, and irregular
pattern on sacral region; dorsolateral stripe-like pattern formed by dark spots from shoulder to groin, with series of
smaller dark spots ventrolaterally on side of trunk; discontinuous black band below canthus rostralis surrounding
nostril; light brown stripe from between nostrils to anteriormost edge of upper jaw; upper lip barred with dark and
light brown; white spots on region posteroventral to eye and on rictus conspicuous; ventral edge of supratympanic
fold and tympanum except for lower part dark brown; groin with dark brown spot extending onto thigh; limbs light
brown dorsally marked with alternating light and dark brown crossbars; basic colouration on upper arm lighter than
on lower arm; dark crossbars restricted to lower arm and fingers; dark brown crossbars on tibia incomplete, absent
on dorsal surface; area of tibio-tarsal articulation not cream-coloured dorsally; throat and chest cream-coloured
with fine, light brown mottling; posterolateral portion of throat without paired unpigmented areas; chin with dark
brown mottling, denser along edge of lower jaw; abdomen cream-coloured with dark brown speckling, forming
diffuse spots of various size; limbs ventrally cream-coloured, heavily dusted with brown on lower arm and
hindlimb; posterior surface of thigh brown speckled with cream; iris red in upper half and along outer margin of
pupil, silver with black reticulation in lower half. In alcohol, colour generally darker, although brown marking on
upper part of tympanum lighter; pattern generally faded but still distinct (Figure 3).
Variation. Individuals of the type series are generally similar to each other in morphology. Variation in size
and body proportions is given in Table 2. Angle of supratympanic fold varies, and in six of eight paratypes, is more
strongly angular than in the holotype. Pectoral glands are more distinct in one paratype. Dark brown marking on
abdomen varies from scattered dots to large round spots. Male specimens from Kinabalu and Crocker do not differ
in size and proportions of body. The single female available is larger (SVL = 35.0 mm) than the largest male (27.9
mm), and has smaller values than males in HL, HW, IND, IOD, UEW, SL, EL, LAL, FLL, TL, FL, and HLL, all
relative to SVL.
TABLE 2 . Measurements in adults of Leptolalax sabahmontanus sp. nov. SVL (MeanцSD, in mm) and medians of
ratios (R) of other characters to SVL, followed by ranges in parenthesis. See text for character abbreviations.
14 males 1 female 3 males
SVL 26.2±0.70 (25.4–27.7) 35.0 (–) 27.1±0.74 (26.5–27.9)
RHL 39.6 (38.2–42.9) 36.3 (–) 38.7 (38.4–41.1)
RHW 34.2 (32.6–36.9) 31.7 (–) 34.3 (33.6–34.4)
RIND 9.4 (8.8–10.5) 8.0 (–) 9.4 (9.3–9.7)
RIOD 11.4 (9.9–12.2) 9.1 (–) 10.1 (9.7–10.6)
RUEW 12.1(11.0–13.0) 10.3 (–) 12.0 (11.6–12.5)
RSL 15.2 (13.2–16.3) 13.1 (–) 15.1 (13.8–15.4)
REL 15.7 (14.4–16.3) 13.7 (–) 15.1 (14.2–16.2)
RN-EL 8.1 (7.3–9.1) – 8.2 (7.9–9.0)
RTD 7.9 (6.9–10.1) 7.1 (–) 7.9 (7.9–8.6)
RE-TL 3.0 (2.0–3.9) 2.9 (–) 3.0 (3.0–4.3)
RLAL 55.2 (51.6–58.5) 50.6 (–) 53.7 (52.0–56.2)
RFLL 73.6 (69.3–79.5) 67.1 (–) 73.4 (72.0–74.7)
RIPL 7.3 (5.4–8.7) 6.0 (–) 7.5 (7.1–7.9)
RTL 54.0 (50.4–56.5) 48.6 (–) 54.1 (52.2–55.8)
RFL 46.5 (42.7–51.2) 42.3 (–) 47.4 (47.3–52.1)
RHLL 170.3 (162.2–186.0) 160.6 (–) 173.9 (169.0–178.9)
RIMTL 5.1 (4.7–6.2) 6.0 (–) 6.0 (4.9–7.5)
R1TOEL 8.2 (7.1–9.0) 7.1 (–) 8.8 (8.7–9.0)
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FIGURE 4. Ventral view of right hand (A) and foot (B) of male holotype of Leptolalax sabahmontanus sp. nov.
(BORNEENSIS 22684). Scale bar = 5 mm.
Call characterics. The advertisement call of four paratypes (BORNEENSIS 22690, 22691; KUHE 48015,
48016) from KNP, recorded at the air temperatures of 17.2–17.7oC were 0.02 to 11.6 (mean±SD = 1.8±2.1, n = 54)
s in duration, with a call interval of 0.1–4.0 (0.5±0.6, n = 48) s (Figure 5, Table 3). A call consisted of a series of 1
to 149 (mean±SD = 24.2±26.9, n = 54) distinct notes. Each note lasted 3–23 (11.7±4.6, n = 46) ms with a note
interval of 54–78 (65.4±6.0, n = 32) ms, and a note repetition rate of 12.5–17.3 (13.4±0.9, n = 51) notes per second.
Each note was composed of 1–2 (1.5±0.5, n = 46) indistinct pulse groups. There was no marked frequency
modulation within a note, and the dominant frequency at the beginning of the call, 6.90–7.35 (7.10±0.11, n = 46)
kHz, ended at 6.75–7.10 (6.94±0.09) kHz, with the decrease of only 0.0–0.4 (0.16±0.08, n = 46) kHz.
The advertisement calls of two males from CRNP (BORNEENSIS 8673 from Trail 11 and 12454 from Mahua)
recorded at the air temperature of 19.8 o
C and 21.3oC, respectively, were basically similar to Kinabalu calls.
However, calls of BORNEENSIS 12454 consisted of longer notes (22–25 [24.0±1.1, n = 11] ms) with shorter
intervals (39–44 [41.9±1.4, n = 10] ms) and higher frequency (initial dominant 7.40–7.60 [7.53±0.06, n = 11] kHz
and final dominant 7.20–7.35 [7.29±0.05] kHz) than in others. Additionally, a harmonic frequency was detected at
20.25–20.45 (20.33±0.08, n = 11) kHz in the calls of BORNEENSIS 12454, probably because of the quieter
environment in which the recording was made.
Comparisons. Leptolalax sabahmontanus differs from congeneric species occurring outside Borneo in the
same manner as in L. fritinniens (Dehling & Matsui 2013), and comparisons with Bornean species are pertinent.
Leptolalax sabahmontanus morphologically differs from all the described Bornean species in the following way.
By the presence of ventral markings, it resembles L. dringi, L. gracilis and L. fritinniens. Of these, the new species
is most similar to L. dringi, but results of statistical tests in males indicated that the new species (Kinabalu and
Crocker combined, n = 17) is smaller than L. dringi from Mulu (n = 7) in SVL (mean = 26.4 mm vs. 29.1 mm in L.
dringi), and that it has smaller eye-tympanum distance (median = 3.0% vs. 4.1%) and inner metatarsal tubercle
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NEW MONTANE LEPTOLALAX FROM BORNEO
(5.1% vs. 5.8%), and larger interorbital distance (11.3% vs. 9.9%), upper eyelid width (12.1% vs. 11.2%), snout
length (15.2% vs. 13.1%), eye length (15.6% vs. 14.0%), and forearm and hand length (55.1% vs. 52.4%), all
relative to SVL, than L. dringi (Mann-Whitney U-test, p < 0.05). Leptolalax sabahmontanus is similar to L. gracilis
in the absence of dark cross bars on the upper arm, but differs from it by a smaller (male SVL 25.4–27.9 mm vs.
31.1–39.2 mm in L. gracilis) and stockier body, and less developed ventral dark marking (body more slender and
dark marking usually conspicuous in L. gracilis). Leptolalax sabahmontanus and L. gracilis are markedly different
in advertisement calls (see below). From L. fritinniens, the new species differs by the smaller body size (male SVL
25.4–27.9 mm SVL vs. 27.4–34.8 mm SVL in L. fritinniens) and having a single, medially arranged vocal sac in
males (vs. bipartite vocal sac in L. fritinniens).
TABLE 3 . Parameters of the advertisement call of six males of Leptolalax sabahmontanus sp. nov., recorded at
Kinabalu National Park (KNP) and Crocker Range National Park (CRNP). Parameter values given as mean ± SD,
followed by sample size (n) and range in parentheses. AT: air temperature; Call L: call length; Call ITV: call interval;
Note RR: note repetition rate; Note L: note length; Note ITV: note interval; Df at beginning: dominant frequency at the
beginning of call; Df at end: dominant frequency at the end of call; Df decrease: decrease in dominant frequency in a call.
The presence of ventral markings clearly differentiates L. sabahmontanus from L. pictus, L. hamidi, and L.
arayai. Furthermore, from L. pictus, the new species differs by having a smaller body size (male SVL 25.4–27.9
mm vs. 31.1–34.0 mm SVL in L. pictus), olive brown dorsum with black pattern without distinct light outlines (vs.
dorsum pinkish brown with dark brown markings with conspicuous thin light outlines in L. pictus). From L.
hamidi, L. sabahmontanus also differs by having a smaller body size (male SVL 25.4–27.9 mm vs. 28.4–28.8 mm
in L. hamidi) and small dorsal markings without light outlines (vs. large dark brown dorsal markings with light
outlines in L. hamidi). From L. arayai, L. sabahmontanus clearly differs by having a smaller body size (male SVL
25.4–27.9 mm vs. 29.2–32.0 mm in L. arayai), not strongly shagreened back, presence of dorsal markings, usually
white venter, and absence of flush colour on ventral side of legs and groin (vs. dorsum tuberculated, without
conspicuous markings, venter yellow, and groin and ventral sides of limbs orange in L. arayai). Finally, the new
species is similar to L. maurus in body size (male SVL 25.4–27.9 mm vs. 25.0–26.9 mm SVL in L. maurus), but
Loc KNP KNP KNP KNP CRNP Trail 11 CRNP Mahua
AT ( oC) 17.2 17.7 17.7 17.7 19.8 21.3
Call L (s) 1.3±0.7 (17) 4.1±4.1 (9) 1.4±1.0 (14) 1.4±1.2 (15) 2.2±1.4 (11) 1.1±0.7 (15)
(0.2–2.6) (0.8–11.6) (0.02–3.8) (0.02–4.0) (0.3–5.0) (0.1–2.4)
Call ITV (s) 0.5±0.4 (16) 0.8±1.4 (8) 0.3±0.2 (13) 0.4±0.1 (13) 0.3±0.1 (10) 0.3±0.1 (14)
(0.2–2.0) (0.2–4.0) (0.1–0.6) (0.1–0.6) (0.2–0.4) (0.1–0.5)
Notes/Call 17.3±8.7 (17) 54.0±53.1 (9) 18.6±12.8 (14) 19.0±15.0 (15) 32.0±20.7 (11) 18.2±10.4 (15)
(3–33) (10–149) (1–48) (1–53) (5–72) (2–37)
Note RR (/s) 13.3±0.8 (17) 13.2±0.2 (9) 13.3±1.0 (14) 13.8±1.2 (15) 14.8±0.9 (11) 16.4±1.0 (15)
(12.7–16.2) (12.8–13.4) (12.5–16.4) (13.0–17.3) (14.2–17.2) (15.6–18.9)
Note L (ms) 6.0±1.3 (11) 18.2±2.4 (11) 10.3±1.0 (13) 12.6±1.2 (11) 15.0±3.0 (10) 24.0±1.1 (11)
(3–8) (14–23) (9–12) (11–15) (10–18) (22–25)
Note ITV (ms) 73.1±1.4 (10) 58.7±2.2 (10) 71.9±2.5 (12) 64.3±1.5 (10) 54.3±2.6 (9) 41.9±1.4 (10)
(71–75) (54–61) (68–78) (62–66) (51–59) (39–44)
Df at beginning
7.02±0.06 (11) 7.15±0.05 (11) 7.23±0.07 (13) 6.99±0.05 (11) 6.80±0.04 (10) 7.53±0.06 (11)
(6.95–7.10) (7.10–7.20) (7.10–7.35) (6.95–7.10) (6.75–6.85) (7.40–7.60)
Df at end (kHz) 6.90±0.07 (11) 7.03±0.03 (11) 7.01±0.04 (13) 6.83±0.04 (11) 6.75±0.05 (10) 7.29±0.05 (11)
(6.80–7.00) (7.00–7.10) (6.95–7.05) (6.75–6.90) (6.70–6.85) (7.20–7.35)
0.13±0.07 (11) 0.13±0.06 (11) 0.22±0.08 (13) 0.16±0.07 (11) 0.05±0.04 (10) 0.24±0.07 (11)
(0.05–0.20) (0.05–0.25) (0.10–0.40) (0.10–0.35) (0–0.10) (0.10–0.35)
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448 · Zootaxa 3753 (5) © 2014 Magnolia Press
differs from it by having an olive brown dorsum with black pattern, and maculated light-colored ventrum (vs.
having a dark brown to black dorsum and ventrum in L. maurus).
FIGURE 5. Sonagram (A, C, E) and wave form (B, D, F) showing advertisement calls of a paratype of Leptolalax
sabahmontanus sp. nov. (KUHE 48016) from the type locality, Silau Silau trail along Sg. Silau-Silau, Kinabalu National Park,
recorded at an air temperature of 17.2°C.
The new species also differs from the known Bornean members acoustically. By lacking clear frequency
modulation within a note, it differs from L. fritinniens, L. hamidi, and L. pictus, although the dominant frequency of
the call of the new species (6.90–7.35 kHz at 17.2–17.7oC) overlaps (7.25–9.20 kHz at 24.3–24.9°C in L.
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NEW MONTANE LEPTOLALAX FROM BORNEO
fritinniens, 6.70–7.30 kHz at 22.9–24.1°C in L. hamidi, and 6.80–7.15 kHz at 19–22°C in L. pictus). The new
species is similar to other species without conspicuous frequency modulation, but the dominant frequency (6.90–
7.35 kHz) of the new species is much higher (6.05–6.40 kHz at unknown temperature in L. dringi; 5.40–5.90 kHz
at 17.4°C in L. arayai; 5.15 kHz at unknown temperature in L. maurus; 2.60–2.80 kHz at 20.0–26.2°C in L.
Range. The species appears to be restricted to Sabah, and has been found around the Headquarters (HQ) of
KNP (Silau Silau trail, Pandanus trail, Bundu Tuhan view trail, and along Sg. Liwagu), Ranau District, and Sg.
Wario, Sayap, Kota Belud District. It also occurs in CRNP, Tambunan District (Trails 4, 5, and 11 of UMS 2002
expedition to the Crocker Range, Ulu Kimanis, and along Sg. Mahua, Mahua). The known localities are all in
mountain regions, ranging in altitude from 754 m to 1500 m a.s.l.
Natural history. Specimens for which data are available were caught in primary rain forest at night 2–50
(median = 5) m away from banks of streams with width of 1–12 (median = 5.5) m. Of the 29 frogs recorded, 20
were found on leaves of short grasses, two on stones at stream banks, two on the trail, three among litters on the
forest floor, and two hidden under rotten log or among root of grasses. The height of the grass at which frogs
perched is 10–100 (median = 30) cm from above the ground. Reproductive details are unknown, but males were
actively calling from late July to mid-August and in mid-December, and these periods must be within the breeding
The new species is found sympatric with L. arayai around Headquarters of Kinabalu National Park (KNPHQ),
L. pictus at Sayap, KNP, and Mahua, CRNP, and L. fritinniens at Trails of Ulu Kimanis, CRNP. The associated
anuran species observed were: Leptobrachium cf. montanum Fischer (at KNPHQ), Leptobrachium cf. abbotti
(Cochran) (at Ulu Kimanis), Leptobrachella baluensis Smith (Ulu Kimanis), Xenophrys dringi (Inger, Stuebing &
Tan) (Ulu Kimanis), Megophrys nasuta (Schlegel) (Mahua), Leptophryne borbonica (Tshudi) (Ulu Kimanis),
Ansonia hanitschi Inger (KNPHQ), A. platysoma Inger (Sayap, Ulu Kimanis, Mahua), A. longidigita Inger (Sayap,
Ulu Kimanis, Mahua), Hylarana raniceps (Peters) (Ulu Kimanis), Staurois parvus Inger & Haile (Ulu Kimanis), S.
guttatus (Günther) (Ulu Kimanis, Mahua), S. latopalmatus (Boulenger) (Ulu Kimanis), S. tuberilingus Boulenger
(KNPHQ, Sayap, Ulu Kimanis), Meristogenys dyscritus Shimada, Matsui, Yambun & Sudin (Sayap), M.
orphnocnemis (Matsui) (Ulu Kimanis, Mahua), M. whiteheadi (Boulenger) (Sayap), M. stigmachilus Shimada,
Matsui, Yambun & Sudin (Mahua), M. kinabaluensis (Inger) (KNPHQ, Ulu Kimanis, Mahua), Huia cavitympanum
(Boulenger) (Sayap), Occidozyga baluensis (Boulenger) (Ulu Kimanis), Ingerana baluensis (Boulenger) (Ulu
Kimanis), Limnonectes palavanensis (Boulenger) (Ulu Kimanis), L. cf. kuhlii (Tschudi) (KNPHQ, Sayap, Ulu
Kimanis), Fejervarya limnocharis (Gravenhorst) (Mahua), Nyctixalus pictus (Peters) (KNPHQ), Philautus
mjobergi Smith (KNPHQ, Sayap, Ulu Kimanis), Ph. petersi (Boulenger) (Sayap, Ulu Kimanis, Mahua), Ph.
aurantium Inger (Sayap, Ulu Kimanis, Mahua), Ph. bunitus Inger, Stuebing & Tan (Sayap, Ulu Kimanis), Ph.
erythrophthalmus Stuebing & Wong (Ulu Kimanis), Ph. macroscelis (Boulenger) (KNPHQ, Ulu Kimanis),
Rhacophorus angulirostris Ahl (KNPHQ, Ulu Kimanis, Mahua), Polypedates leucomystax (Gravenhorst) (Mahua),
P. otilophus (Boulenger) (Ulu Kimanis), Kalophrynus heterochirus Boulenger (Ulu Kimanis), K. baluensis Kiew
(KNPHQ), K. pleurostigma Tschudi (Crocker Trails), and Chaperina fusca Mocquard (Sayap, Ulu Kimanis,
Malkmus et al. (2002) described the call characteristics of L. dringi from the Kinabalu region, but these do not
match the spectrogram displayed (Malkmus et al. 2002: Figure XLVIII). Matsui & Dehling (2012) suspected that
the spectrogram shown in Malkmus et al. (2002) is from Dring’s original recorded tape, and that the description of
the call is based on the call of another species of Leptolalax from Kinabalu region. The call characteristics of L.
dringi described in Malkmus et al. (2002: 101) also differ from those analyzed by us. The call length (1.8–3.5 s in
Malkmus et al.’s L. dringi vs. 0.02 to 11.6 s in our calls), number of notes in a call (8–10 vs. 1–149), the note
interval (60 ms vs. 54–78 ms), and the dominant frequency (7.0–7.5 kHz vs. 6.9–7.4 kHz) overlap between the two
studies, but the note length is longer (35–40 ms vs. 3–23 ms) and the note repetition rate is lower (11.0–11.5 per s
vs. 12.5–17.3 per s) in the call described by Malkmus et al. (2002: 101) than in the call we analyzed. Malkmus et
al. (2002) also noted the presence of distinct frequency modulation in the call of L. dringi from Kinabalu, but
MATSUI ET AL.
450 · Zootaxa 3753 (5) © 2014 Magnolia Press
frequency modulation was not conspicuous in our calls. From the description of call characteristics, we suspect that
L. dringi described in Malkmus et al. (2002) may represent another species, most probably, L. fritinniens, recently
described from Sarawak, whose calls are characterized by conspicuous frequency modulation (Dehling & Matsui
2013), and which is very common at lower elevations of Kinabalu, especially around Poring, which was listed as a
locality of L. dringi by Malkmus et al. (2002).
Besides confusions with L. dringi, L. sabahmontanus has been identified as L. gracilis in many museum
collections, as is the case in L. fritinniens (Dehling & Matsui 2013). These four species are superficially similar in
the possession of ventral marking despite very different acoustic characteristics. As a result of the description of L.
sabahmontanus as a new species, the distribution range of L. dringi is restricted to the type locality of high altitudes
(ca. 1800 m a.s.l.) on Gunung Mulu in Sarawak. Between known localities of L. sabahmontanus (southernmost
locality at Ulu Kimanis of the Crocker range) and L. dringi (Gunung Mulu), there are few mountains over 754 m in
altitude (the known lower limit of L. sabahmontanus), and no records of either species are available. Instead, L.
pictus and L. fritinniens are widely distributed there (Matsui, unpublished data).
The new species has been found syntopically with other species of Leptolalax in all known localities. On Mt.
Kinabalu, it occurs syntopically with L. arayai around the Park Headquarters (Silau Silau, Pandanus, and Liwagu
trails), and seems to be much more abundant than L. arayai. Another congener, L. maurus seems to occur at higher
elevations than the new species, and at present there is no record of their coexistence. Leptolalax sabhmontanus
occurs almost syntopically with L. pictus at Sayap field station, but is much less in number than the latter species.
On the Crocker range, the new species is also syntopic with L. fritinniens at Trail 4, Trail 5, and Trail 11, and L.
pictus at Mahua, but is much less frequently encountered than the syntopic congeners in all these localities.
Coexistence of more than one species of Leptolalax at one locality seems to be not uncommon at least around
Kinabalu and Crocker ranges of north-western Borneo, as shown above. On Mt. Kinabalu, L. sabhmontanus co-
occurs L. pictus or L. arayai, both with larger body size, but not with L. maurus with similar body size. On the
Crocker range, L. sabhmontanus co-occurs with larger-sized L. fritinniens and L. pictus. Most probably, differences
in body size allows co-existence through resource partitioning, i.e., segregation in feeding behavior and
microhabitat selection. Additionally, body size difference, together with differential acoustic characteristics, would
prohibit interspecific hybridization.
Matsui et al. (2010) studied molecular phylogeny in Southeast Asian stream-adapted toads of the genus
Ansonia and recognized two major clades, which tend to differ in body size: species of one clade are small-sized
while those of another clade are medium- to large-sized. They reported that syntopic distribution of two species is
commonly observed in Bornean Ansonia, and species belonging to different major clades usually coinhabit in such
cases. From these observations, Matsui et al. (2010) surmised that differential body size seems to be related to the
prevailing sympatric distribution of two Ansonia species in many localities of Borneo.
In the case of Bornean Leptolalax, detailed phylogenetic relationships are not yet clarified, but limited data
shown in this study (Figure 1) indicate that the body size is not directly related to phylogeny because large-sized L
arayai and small-sized L. maurus, and large-sized L. pictus/L. fritinniens and small-sized L. dringi/L.
sabahmontanus, respectively, showed sister group relationships. Thus, although body size in Bornean Leptolalax
might have been affected by resource allocation related to sympatric distribution, accumulation of more data on the
relationships between detailed distribution and body size, and application of pertinent analyses to them are
necessary in order to confirm this hypothesis.
The Economic-Planning Unit of Malaysia (EPU: 40/200/19 SJ. 1158) and Sabah Parks (SP) kindly permitted M.
Matsui to conduct the project, and UMS, Japan International Cooperation Agency (JICA), and SP kindly provided
all the facilities for conducting research. M. Matsui is grateful to the following for their encouragements and/or
permission to conduct research and field companionship: H. Akiyama, L. Apin, Y. Hashimoto, J. K. B. Kueh, T.
Kusano, M. Maryati, T. Shimada, A. Sudin, T. Tachi, and S. Yasuma. M. Matsui also thanks K. Eto and A. Hamidy
for help in the laboratory, and N. Jamili and M. B. Lakim (SP), and M. Maryati (UMS) for allowing him to examine
specimens under their care. Field trips and examination of museum specimens were made possible by Grants under
The Monbusho /JSPS Scientific Research Programs (Nos. 10041166, 15370038, 20405013, and 23405014 to MM
and 17405018 to KN).
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APPENDIX. Comparative material examined (all from Borneo).
Leptolalax arayai: SP 01804, HQ of KNP, Sabah (holotype); BORNEENSIS 22624, 22692, 22693, 22701 Silau Silau, KNP,
Sabah; BORNEENSIS 22739–22747 Pandanus Trail, KNP, Sabah; BORNEENSIS 22903, 22931, Mesilau, KNP, Sabah.
Leptolalax dringi: KUHE 55610, 55612, 55613, Sg. Tapin, Mulu NP, Sarawak.
Leptolalax fritinniens: KUHE 10534 (paratype), 53676 (paratype), 53678–53681 (paratypes), Camp 5 of Mulu NP, Sarawak;
KUHE 55371, Base Camp of Mulu NP, Sarawak; KUHE 55625, Camp 1 of Mulu NP, Sarawak; BORNEENSIS 23380–
23382, 23384, 23386–23388, 23390–23392, 23414, Sg. Kilanpug of CRNP, Sabah; BORNEENSIS 4416, 12805–12807,
12814, 12816, 12912, 12913, 12915, Ulu Senagan, CRNP, Sabah; BORNEENSIS 8710, 8712, 8714, 8804, 8828–8831,
Trail 4 of Ulu Kimanis, CRNP, Sabah; BORNEENSIS 8180, 8198, 8400, 12878, Trail 5 of Ulu Kimanis, CRNP, Sabah;
BORNEENSIS 8497, 8498, KUHE 39205, Trail 11 of Ulu Kimanis, CRNP, Sabah; BORNEENSIS 8715, 12731, 12747–
12750, 12784–12787 TBC of Ulu Kimanis, CRNP, Sabah; BORNEENSIS 12843–12847 Masak, CRNP, Sabah;
BORNEENSIS 12635, 12637, Mahua, CRNP, Sabah; BORNEENSIS 22436, Tamparuli, Sabah; BORNEENSIS 22460,
Bundu Tuhan, Sabah; BORNEENSIS 2049, 6001–6003, Serinsim Substation of KNP, Kota Marudu, Sabah;
BORNEENSIS 2487, 22831–22836, 22866–22868, KUHE 39302–39305 Poring, KNP, Sabah; BORNEENSIS 22335,
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22336, 22486–22488, Paginatan, Sabah; BORNEENSIS 4051, 9112. 9173, 9199–9203, 9209, 9248–9250, 22060, 22061,
22093, 22094, 22148, Tawau Hills NP, Sabah; BORNEENSIS 22311, Tabin WLS, Sabah.
Leptolalax gracilis: BMNH 188.8.131.52 = 1947.2.25 (holotype), KUHE 10634-10637, 10717, 12019, 12020, 12023, 17273,
17183, 17665, 19567, 33948, 42644, 42645, 42663, 53042, 53043, 53164, 53173, 53239, 53771, 53777, 53782, 53788.
53791, 53809, 54538–54541, 55382, 55388, 55390, 55395–55398, 55400–55403, 55439, 55440, Gunung Serapi, Kubah
NP, Sarawak; KUHE 17133, 17136, 17168, 17183, Gading NP, Sarawak; KUHE 53230, 53231, Serian, Sarawak; KUHE
53689, 53690, Bukit Kana, Sarawak; KUHE 53889, 53921, 53922, 54419, 54431, 55464–55466, 55470, 55476–55478,
Gunung Penrissen, Sarawak; KUHE 55418, 55425, Santubong, Sarawak; KUHE 55336, 55624, Camp 1 of Mulu NP,
Leptolalax hamidi: KUHE 17374 (paratype), 17545 (paratype), 17546 (holotype), 17547, Bukit Lanjak, Sarawak.
Leptolalax maurus: SP 26749, 26750, 26756, 26757, KNP, Sabah.
Leptolalax pictus: BORNEENSIS 8871–8876, 12422, 12426, 12501–12503, 12547, 12549, 12552, 12600–12602, 12604,
12606, 12697, Mahua, CRNP, Sabah; BORNEENSIS 4408, Tambunan, Sabah; BORNEENSIS 2322, 2349, 2350, 2355,
2401, 2430, 2435, Sayap, KNP, Sabah; BORNEENSIS 22350, 22351, 22459, KUHE 39393, 39394, Bundu Tuhan, Sabah;
BORNEENSIS 22837–22840, 22859, 22864, 22894, KUHE 14529, 39297, 39298, Poring, KNP, Sabah; KUHE 11226,
53099–53102, Bario, Sarawak.