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Zootaxa 2814: 33–49 (2011) ISSN 1175-5326 (print edition)

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Copyright © 2011 · Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)

Taxonomic revision of one of the Old World’s smallest frogs, with description
of a new Bornean Microhyla (Amphibia, Microhylidae)

MASAFUMI MATSUI
Graduate School of Human and Environmental Studies, Kyoto University, Kyoto 606-8501, Japan.
E-mail: fumi@zoo.zool.kyoto-u.ac.jp

Abstract

An examination of the holotype of Microhyla borneensis Parker 1928 revealed that this nomen applies to a miniaturized
narrow-mouthed frog from Borneo that recently was described as Microhyla nepenthicola Das & Haas 2010. This is con-
firmed concordantly by body size of the female holotype of M. borneensis, and by its reduced extent of toe webbing and
rather rounded, short snout profile. Consequently, the name M. nepenthicola is to be considered as a junior synonym of
M. borneensis, and an old report of nepenthiphilous breeding habits of M. borneensis is therefore justified. A sympatric
larger species usually treated as M. borneensis has no scientific name and is described here as M. malang sp. nov. The
new species and M. borneensis are sister to each other, and together are closely related to M. mantheyi from Peninsular
Malaysia. These three species are morphologically very similar, but can be distinguished by body size, color pattern, and
extent of toe webbing. Phylogenetic relationships, miniaturization, and larval oral morphology in the genus Microhyla are
discussed.

Key words: Microhyla, new species, synonymy, mitochondrial DNA, phylogeny, body size, Borneo, miniaturization

Introduction

The genus Microhyla Tschudi has a wide range of distribution from the Ryukyu Archipelago of Japan through Tai-
wan, southern China, Southeast Asia, and much of South Asia including Sri Lanka, and encompasses about 30 spe-
cies (Frost 2010). Within Southeast Asia, Borneo is renowned for its high diversity of amphibian species (Inger &
Tan 1996; Matsui 2006), but diversity of Microhyla on the island is not high, with only five species (M. berdmorei
[Blyth], M. borneensis Parker, M. perparva Inger and Frogner, M. petrigena Inger and Frogner, and M. maculifera
Inger) recorded over the past two decades (Frost 2010).
Das & Haas (2010) recently described M. nepenthicola from Sarawak, emphasizing its small size as the Old
World’s smallest frog, and recorded its unique nepenthiphilous breeding habits as their original finding. In the
description of M. nepenthicola they treated a sympatric, and sometimes syntopic, larger sized species as M.
borneensis. Recognition of the larger species as M. borneensis by these authors began when Das et al. (2007a) sep-
arated the Malay Peninsula population (Berry 1975; Chan-ard 2003) from the Bornean population as a distinct spe-
cies M. mantheyi, and Das & Haas (2007b) reviewed the amphibian fauna of the Matang Range, Sarawak, and
briefly introduced M. nepenthicola as an undescribed species.
Parker (1934) identified a nepenthiphilous larva from Kuching, near the type locality of M. borneensis, Bidi
(Kidi in the original description), as that species. Inger (1966), however, doubted the species identification and
assigned the unique larvae to another microhylid genus Kalophrynus. Inger (1966) instead described larvae from
isolated pools of Third and Fourth Divisions of Sarawak as M. borneensis and most subsequent authors (e.g. Malk-
mus et al. 2002; Das et al. 2007a, b) followed this view.
During herpetological fieldwork in Sarawak, I have been studying nepenthiphilous larvae and their parental
species at various altitudes of Gunung (= Mt.) Serapi, including the type locality of M. nepenthicola (Das & Haas
2010). The species is identified as M. borneensis by Inger's (1966) key, except for definitely smaller adult body
size. At the same locality, I found specimens of Microhyla sympatric with the small form (Fig. 4). They also keyed

Accepted by M. Vences: 14 Mar. 2011; published: 11 Apr. 2011 33


out to M. borneensis according to Inger (1966), and their body size better fits his description than the small form. I
have also detected the large form and larvae assigned to it in ponds and stream-side pools in Sarawak and Sabah.
Although both forms could be identified as M. borneensis by Parker's (1934) monograph, the original description
of the species (Parker 1928) led me to consider the small, nepenthiphilous form as true M. borneensis. These were
confirmed in a subsequent examination of the holotype of M. borneensis.
These collective observations suggest that the taxonomic treatment of Das & Haas (2010) should be re-exam-
ined. At the same time, the nepenthiphilous breeding habits of M. borneensis as originally reported by Parker
(1934) should be considered valid. In the present paper, I will show the results of my own examination of the small
form (topotypic M. nepenthicola) and the sympatric large form (M. borneensis sensu Das & Haas), as well as of the
holotype of M. borneensis stored in BM. Based on these results, I will elucidate the correct taxonomic status of the
two forms and describe the large form as a new species.

Material and methods

Fieldwork was conducted between 1979 and 2010 in Sarawak (Gunung Serapi, now partly in Kubah National Park,
Matang Range, and Kanowit) and Sabah (Poring on Mt. Kinabalu, Sepilok near Sandakan, Tawau Hills National
Park, and Tabin Wildlife Reserve). In the field, I recorded calls using a cassette tape recorder (Sony TC-D5) with
an external microphone (Sony ECM-23) or a digital recorder (Zoom H2). I fixed specimens collected in 10% form-
alin, later preserved in 70% ethanol, and stored at the Graduate School of Human and Environmental Studies,
Kyoto University (KUHE), the Sarawak Museum (SM), and the Institute for Tropical Biology and Conservation,
University Malaysia Sabah (BORNEENSIS). Since 1989, I have also taken tissues from some specimens for later
phylogenetic analyses.
For adult specimens, I took the following 19 body measurements to the nearest 0.05 mm with dial calipers, fol-
lowing Matsui (1984) and Matsui (1994) : (1) snout-vent length (SVL); (2) head length (HL), from tip of snout to
hind border of angle of jaw (not measured parallel with the median line); (3) snout length (SL); (4) eye length (EL);
(5) nostril-eyelid length (N-EL); (6) head width (HW); (7) internarial distance (IND); (8) interorbital distance
(IOD); (9) upper eyelid width (UEW); (10) forelimb length (FLL); (11) lower arm and hand length (LAL), from
elbow to tip of third finger; (12) first finger length (1FL), from distal end of inner palmar tubercle to tip; (13) hand
length (HAL), from proximal end of outer palmar tubercle to tip of third finger; (14) inner palmar tubercle length
(IPTL); (15) outer palmar tubercle length (OPTL); (16) hindlimb length (HLL); (17) tibia length (TL); (18) foot
length (FL); and (19) inner metatarsal tubercle length (IMTL). Additionally, I also measured: (20) first toe length
(1TOEL), from distal end of inner metatarsal tubercle to tip of first toe; (21) outer metatarsal tubercle length
(OMTL); (22) first finger width (1FW), measured at the distal phalanx; (23–25) finger disk diameters (2-4FDW);
(26–30) toe disk diameters (1–5TDW); and (31–36) finger lengths (1–3FLO, 2–4FLI), for outer side (O) of the first,
inner side (I) of the fourth, and both sides of the remaining fingers, measured between tip and the junction of the
neighboring finger. I made all measurements under a binocular dissecting microscope. I followed the system of
description of toe-webbing states used by Savage (1975).
Examination of the holotype of M. borneensis housed in BM was made on 20 October 1998. For comparisons,
I examined museum specimens, including some types, of many described taxa of the genus Microhyla (see Appen-
dix 1).
For preserved larvae, I took the following 13 measurements to the nearest 0.01 mm using a binocular dissect-
ing microscope equipped with a micrometer: (1) total length (TOTL); (2) head-body length (HBL); (3) maximum
head-body width (HBW); (4) maximum head-body depth (HBD); (5) eye-snout distance; (6) eyeball diameter; (7)
interorbital distance; (8) snout-spiracle opening distance; (9) oral disk width; (10) tail length; (11) maximum tail
depth; (12) maximum tail width, and (13) maximum tail muscle depth. Staging followed Gosner's (1960) table.
I analyzed the recorded calls using the computer programs SoundEdit Vers. 2 and SoundEdit Pro (MacroMind-
Paracomp, Inc.) on a Macintosh computer. Methods of analyses are as described elsewhere (Matsui 1997).
For genetic comparisons, I used DNA sequences of the large type (M. borneensis sensu Das & Haas, 2010) and
its putative larva from a pool, the small type (M. nepenthicola of Das & Haas, 2010) and its putative nepenthip-
hilous larva, M. achatina, M. berdmorei, M. butleri, M. heymonsi, and M. mantheyi. For comparisons, DNA
sequences already reported (Matsui et al. 2005) were obtained from GenBank for M. okinavensis (AB201173,
AB201184). As outgroup, I used Calluella minuta, Chaperina fusca, and Micryletta inornata (Table 1). Methods

34 · Zootaxa 2814 © 2011 Magnolia Press MATSUI


for phylogenetic analyses follow Matsui et al. (2010b) with slight modification. Total DNA was extracted, ampli-
fied by PCR, and sequenced using the primers 12SA-L (Palumbi et al., 1991), 12SF (Goebel et al., 1999), H1548
(Matsui et al., 2005), 12SL 1 (Matsui et al., 2010a), L1507 (Matsui et al., 2010b) for 12S, and 16H1 and 16L-1
(Hedges, 1994), will6 (Wilkinson et al., 2002), L2188 (Matsui et al., 2006), L1879 and H1923 (Matsui et al.,
2010b) for 16S rRNA. The resultant sequences were deposited in GenBank (AB598295—598342: Table 1). The
alignment matrix with 1735 nucleotide sites (894 sites for 12S rRNA; 841 for16S rRNA) was subjected to estimate
phylogenetic relationships using maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference.
Pairwise comparisons of uncorrected sequence divergences (p-distance) were also calculated. Details for these pro-
cedures are given in Matsui et al. (2010a).

TABLE 1. Samples used for phylogenetic analysis, and GenBank accession numbers. BOR: BORNEENSIS collection, Uni-
versity Malaysia Sabah; KUHE: Kyoto University, Human and Environmental Studies; MZB: Museum Zoologicum Bogo-
riense; SM: Sarawak Museum.
Sample Species Locality Voucher Accession No.
No. 12S 16S
1 Microhyla sp. large Gunung Serapi, Sarawak, Malaysia KUHE 53018 AB598295 AB598319
2 Microhyla sp. large Gunung Serapi, Sarawak, Malaysia KUHE 53034 AB598296 AB598320
3 Microhyla sp. large (Larva) Gunung Serapi, Sarawak, Malaysia KUHE NNL AB598297 AB598321
4 Microhyla sp. large Kanowit, Sarawak, Malaysia KUHE 42597 AB598298 AB598322
5 Microhyla sp. large Tawau Hills, Sabah, Malaysia BOR 9090 AB598299 AB598323
6 Microhyla sp. large Tawau Hills, Sabah, Malaysia BOR 9210 AB598300 AB598324
7 Microhyla sp. large Tawau Hills, Sabah, Malaysia BOR 9211 AB598301 AB598325
8 Microhyla sp. large (Larva) Tawau Hills, Sabah, Malaysia KUHE NNL AB598302 AB598326
9 Microhyla sp. small Gunung Serapi, Sarawak, Malaysia KUHE 53020 AB598303 AB598327
10 Microhyla sp. small Gunung Serapi, Sarawak, Malaysia KUHE 53033 AB598304 AB598328
11 Microhyla sp. small Gunung Serapi, Sarawak, Malaysia KUHE 53165 AB598305 AB598329
12 Microhyla sp. small Gunung Serapi, Sarawak, Malaysia SM (081126) AB598306 AB598330
13 Microhyla sp. small (Larva) Gunung Serapi, Sarawak, Malaysia KUHE NNL AB598307 AB598331
14 Microhyla sp. small (Larva) Gunung Serapi, Sarawak, Malaysia KUHE NNL AB598308 AB598332
15 Microhyla mantheyi Templer Park, Selangor, Malaysia KUHE 15726 AB598309 AB598333
16 Microhyla mantheyi Temerloh, Pahang, Malaysia KUHE 52556 AB598310 AB598334
17 Microhyla achatina Ungaran, Java, Indonesia MZB Amp AB598311 AB598335
16401
18 Microhyla heymonsi Ranong spa, Ranong, Thailand KUHE 23856 AB598312 AB598336
19 Microhyla okinavensis Amamioshima Is., Kagoshima, Japan KUHE 12840 AB201173 AB201184
20 Microhyla berdmorei Gombak, Selangor, Malaysia KUHE 52034 AB598314 AB598338
21 Microhyla butleri Nongten, Vientiane, Laos KUHE 40649 AB598315 AB598339
22 Calluella minuta Temerloh, Pahang, Malaysia KUHE 52463 AB598316 AB598340
23 Micryletta inornata Mae Yom, Phrae, Thailand KUHE 20497 AB598317 AB598341
24 Chaperina fusca Crocker, Sabah, Malaysia BOR 8478 AB598318 AB598342

Results

Morphological analyses

Comparisons of two forms with two species shown in Das & Haas (2010). The small form and the large form of
M. borneensis from Gunung Serapi matched morphologically with the descriptions of M. nepenthicola and M.

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 35
borneensis, respectively, of Das & Haas (2010). Snout-vent lengths of the small form (19 males 10.4–13.4 mm, 5
females 16.7–18.9 mm) and the large form (9 males from Serapi 19.4–21.3 mm) well conformed to those of M.
nepenthicola (8 males 10.6–12.8 mm, 2 females 17.9 and 18.8 mm) and M. borneensis (males to 18 mm, females to
23 mm, sample size not indicated), respectively, given by Das & Haas (2010). This concordance is natural because
they were all collected actually from the same locality. A single adult female of the large form available (KUHE
44228) is from Sabah, and has the SVL of 22.2 mm. Another female, also from Sabah (KUHE 44229) has imma-
ture ovaries, but has the SVL of 18.2 mm. Other than the body size difference, Das & Haas (2010) differentiated
their M. borneensis from M. nepenthicola as follows: the possession of a weak supratympanic fold (vs. absent in M.
nepenthicola); Finger I reduced, but projecting from margin of carpal margin (vs. reduced to a tubercle); tubercles
forming a low ridge on flanks (vs. absent); and a light-edge to the dark pattern on dorsum (vs. absent).
However, all of these diagnostic characteristics, except one, include discrepancies in the description by Das &
Haas (2010). In describing the holotype of M. nepenthicola, these authors repeatedly mentioned the presence of a
weak fold across the supratympanic region (p. 41, line 7), or of a weak and narrow supratympanic fold (p. 43, line
10). In the diagnosis of the species, it is stated that Finger I is reduced to a nub proximal to Finger II in males (p. 40,
line 10 from bottom), but in the section on variation (p. 43, line 12 from bottom), females were noted to show
greater development of Finger I, with one phalange clearly visible. As for tubercles on the flanks, the holotype was
shown to have a dorsum with low tubercles that are relatively more distinct on the flanks (p. 43, line 9). Thus, the
presence or absence of a supratympanic fold, reduction of the first finger, and development of tubercles on the
flanks, listed as characters to differentiate M. borneensis and M. nepenthicola by Das & Haas (2010) are all not val-
idated by themselves. The single characteristic left is the presence in M. borneensis and absence in M. nepenthicola
of a light-edge to the dark pattern on the dorsum.
In my sample, a weak supratympanic fold was always present, though sometimes only traceable, in both the
small and large forms. The first finger was usually reduced in males of the small form, but some had a short, but
distinct phalange like most females. A few females had a reduced first finger, but most had the finger as well devel-
oped as that seen in the large form. The degree of development of tubercles on the flanks was variable, and could
hardly be used to distinguish the two forms. By contrast, the dark pattern on the dorsum was found edged by a light
line in most specimens of the large form. However, it was also clear in a few males of the small form.
In my examination of the two forms, the more reliable characteristic that differentiated them was the toe web-
bing, which was much less developed in the small form: in males, phalanges free of web in the large form were two
(vs. often 21/3 or more in the small form) on the inner (preaxial) side of the second toe, one (vs. often 11/2 and up to
12/3) on the outer (postaxial) side of the second, up to 22/3 (vs. more than three) on the inner side of the third, usually
up to 12/3 (vs. more than two) on the outer side of the third, constantly three or less (vs. more than 31/2 and often
33/4) on both sides of the fourth, and constantly one (vs. two) on the fifth (Fig. 1).
Examination of holotype of Microhyla borneensis and identity of two species. As noted above, I examined
the holotype of M. borneensis in 1998. The holotype of M. borneense (original spelling) collected by Brooks from
Kidi (changed to Bidi in Parker 1934) district, Sarawak, and originally numbered as BM 1911. 1. 30. 43 (Parker
1934), has been renumbered as BM 1947. 2.11.82 as noted by Inger (1985). A slight cut had been made in the past
dorsolaterally on the right side of the holotype, and I could glance ripe oocytes from the slit. Thus the fact that the
holotype is an adult female, as stated in the original description and on the catalogue of the museum, was ascer-
tained. Although the specimen was reported to have a SVL of 18 mm (Parker 1928), it was smaller in my measure-
ments, measuring 17.3 mm (HW 5.7 mm and TL 11.0 mm), probably partly due to shrinkage of the specimen and
partly to different manner of the measurement. In the description of toe web development, Parker (1928) seems to
have mistakenly treated outer margins of the first and second toes as inner ones. The web formula transformed from
the verbal description of Parker (1928) is I 1–? II 1–2 III 2–3 IV 3–2 V, but is I 1–2 II 1–3 III 21/4 –31/3 IV 31/2–2 V,
according to my observation (see Fig. 2D).
Otherwise, the original description (Parker, 1928) was confirmed in all respects, except for color, which has
seriously faded, while the pattern of markings shown in the original description was not clear except for a dark
brown streak on the flanks (Fig. 2). Even so, the body size of the adult female holotype of M. borneensis (SVL < 18
mm) was within the range of the females of the small form (16.7–18.9 mm, see above) and was much smaller than
the female large form (22.2 mm in an adult and 18.2 mm in a juvenile). Furthermore, the toe webbing (I 1–2 II 1–3
III 21/4 –31/3 IV 31/2–2 V ) was much more similar to the small form (I 1–21/3 II 11/2–3 III 2 –31/2 IV 31/2–2V) than to

36 · Zootaxa 2814 © 2011 Magnolia Press MATSUI


the large form (I 1–2 II 1–22/3 III 12/3 –31IV 3–1 V). These two characteristics clearly indicated that the holotype of
M. borneensis and the small form (i.e. M. nepenthicola), rather than the large form (i.e. M. borneensis sensu Das &
Haas [2010]), were conspecific.

FIGURE 1. Ventral views of feet of the large form (A-D: males from Gunung Serapi, Sarawak [A: KUHE 17217; B: KUHE
17245; C: KUHE 17658; D: KUHE 53176], E: female from Poring, Sabah [KUHE 44228]) and the small form of Microhyla
(F-J: males [F: KUHE 10693; G: KUHE 10695; H: KUHE 10743; I: KUHE 53241; J: KUHE 53938]; K-O: females [K: KUHE
12067; L: KUHE 26419; M: KUHE 53020; N: KUHE 53165; O: KUHE 53582], all from Gunung Serapi, Sarawak). Not to
scale.

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 37
FIGURE 2. Female holotype of Microhyla borneensis (BM 1947. 2.11.82); dorsal (A), ventral (B) and lateral views (C), and
ventral view of left foot (D). Scale bar in (A) and (B) = 10 mm.

Molecular phylogenetic analyses

Concatenated fragments of mtDNA genes for 24 samples, including outgroup taxa, were 1701 bp in length. Of
1735 nucleotide sites, 576 were variable, and 347 were parsimoniously informative. The best substitution model
was the general time reversible model (GTR: Rodriguez et al. 1990) with proportion of invariable sites (I) of
0.3961 and a gamma shape parameter (G) of 0.4764. Four equally most parsimonious trees (L = 1357 steps, CI =
0.596, RI = 0.653) were obtained. The likelihood values (-lnLs) of the ML and BI trees were 8148.3828 and
8155.3564, respectively.
Because phylogenetic analyses employing the three different analysis methods yielded same topologies, only
the Bayesian tree is presented in Fig. 3. Samples of Microhyla formed a monophyletic group with respect to the
three outgroup species, but five clades exhibited unresolved relationships. Of these five, M. achatina, M. mantheyi,
and the large and small forms were monophyletic (76, 96, and 100% in MP and ML BS and BPP, respectively).
Within this group, M. mantheyi tended to be a sister species to the clade including samples of the large and small
forms, but their monophyly was not supported by MP BS (63/91/100%). Monophyly of the large and small forms
as well as of each of them was fully supported. In the clade of the large form, each of Sarawak and Sabah samples,
both including their putative larvae, was monophyletic (98, 98, and 100%, and 100, 99, and 100%, respectively),
and clearly split from each other. The genetic distances (uncorrected p-distance) between sympatric larval and adult
samples in each of the two forms were very small, 0.24–1.12% for larvae from ponds and the large form, and 0.01–
0.18% for larvae from Nepenthes pitchers and the small form. The distances between samples of the large form
from Sarawak and Sabah were slightly larger (2.72–3.01 [mean = 2.86]%), but the differences between the two
forms were much greater (4.61–5.27 [mean = 4.89]%). The mean distances of the large and small forms from M.
mantheyi were 7.21–7.69 [mean = 7.46]% and 6.98–7.16 [mean = 7.09]% , respectively.
When my sequences are compared with shorter sequences given by Das & Haas (2010) in GenBank data,
uncorrected p-distances were only 0–0.3% between the small form and M. nepenthicola, and 0.4–0.7% between the
large form and their M. borneensis, and again clearly indicated associations of the small form with M. nepenthicola

38 · Zootaxa 2814 © 2011 Magnolia Press MATSUI


and the large form with M. borneensis sensu Das & Haas (2010). As noted above, the small form actually repre-
sents true M. borneensis, with M. nepenthicola as its synonym, while the large form has no available name.
Because the large form also differs morphologically from all known species of Microhyla, I describe it below as a
new species.

FIGURE 3. Bayesian tree of a 1735 bp sequence of mitochondrial 12S and 16S rRNA for samples of Microhyla (for the sam-
ple details, see Table 1). Numbers above or below branches represent bootstrap supports for MP and ML inferences, and Bayes-
ian posterior probability (MP-BS/ML-BS/BPP).

Microhyla malang sp. nov.

Synonymy: Microhyla borneensis: Parker 1934, p. 128 (part); Inger 1966, p. 151 (part); Inger & Stuebing 1997, p. 108; Malk-
mus et al. 2002, p. 131; Das et al. 2007a, p. 101; Das et al. 2007b, p. 156; Das & Haas, 2010, p. 45.

Holotype. KUHE 17245, an adult male from Gunung Serapi, Matang Range, in the suburbs of Kuching, Kuching
Division, Sarawak, Malaysia (01o36'N, 110o11'E, alt. 290 m a.s.l.; now a part of Kubah National Park), collected on
15 August 1993 by Masafumi Matsui.
Paratypes. Eleven adult males: KUHE 17217, 17236, 17658 from the type locality, collected on 12 August, 12
August, and 9 September, 1993, respectively; SM unnumbered (former KUHE 53034), KUHE 53018 from the type
locality, collected on 14 August 2008; KUHE 53166 , 53176 from the type locality, collected on 26–27 August
2008; Universiti Malaysia Sarawak (UNIMAS) R2110 from the type locality by Ramlah Zainudin; and KUHE
42549, 42550, 42597 from Kanowit, Sibu Division, collected on 20, 20, and 22, respectively, November 2008 by
Daicus M. Berabut, Norhayati Ahmad, Kanto Nishikawa, and Masafumi Matsui.
Referred specimens. Universiti Kebangsaan Malaysia (UKM) HC 00392 an adult male from Lanjak Entimau
Wildlife Sanctuary, Sri Aman Division, Sarawak, collected in 2008 by Daicus M. Berabut and Norhayati Ahmad;
KUHE, two unnumbered batches of tadpoles, from the type locality and a stream near the type locality, collected on
27 August 2010 by Masafumi Matsui and Koshiro Eto; KUHE 44228, an adult female from Poring Hot Spring,

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 39
Pantai Barat Division, Sabah, collected on 21 March 1979 by Masafumi Matsui (Matsui, 1979); KUHE 44229, a
juvenile female from Sepilok, Sandakan Division, Sabah, collected in August 1979 by Ryozo Yoshii; Sabah Park
Collection (SP) 02662 from Imbak, Telupid, Sandakan Division, Sabah, collected on 5 July 2000 by Maklarin
Lakim, Paul Yambun, and others. BORNEENSIS 9090, 9210, 9211 from Tawau Hills National Park, Tawau Divi-
sion, Sabah, collected from 28 July to 1 August 2004 by Masafumi Matsui, Kanto Nishikawa, and Tomohiko Shi-
mada; KUHE, an unnumbered batch of tadpoles, from Tabin Wildlife Reserve, Tawau Division, Sabah, collected
on 3 August 2004 by Tomohiko Shimada and Masafumi Matsui.
Etymology. The specific epithet malang is a Malay word denoting unlucky, alluding to the long history of tax-
onomic confusion in which it was called under the name of its sister species M. borneensis. It is also an allusion of
Matang, the type locality of the species in western Sarawak.
Diagnosis. A medium form of Microhyla, adult males about 18–22 mm in SVL; snout obtusely pointed,
rounded in profile; first finger short, less than one-fourth of third; tips of three outer fingers weakly dilated, form-
ing weak disks, dorsally with median longitudinal groove; outer palmar tubercle single; tibiotarsal articulation
reaching far beyond snout tip; tips of toes distinctly dilated into disks, dorsally with median longitudinal groove;
toes on outer side of second 1, inner side of third 22/3, outer side of third 11/2 – 12/3 , inner and outer sides of fourth
3, and inner side of fifth 1 phalange(s) free of web; inner and outer metatarsal tubercles present; eyelid without
supraciliary spines; top of snout undifferentiated from the brown forehead; vertebral stripe absent; dark dorsal
markings usually narrowly outlined in white; a cream stripe below supratympanic fold, not bordered by black
above; a narrow, broken, black lateral stripe from above arm to half length of trunk; tail of larvae with two black
markings separated by an oblique white marking at middle, and transparent filament at tip.
Description of holotype (measurements in mm). SVL 21.4; habitus stocky, body rounded (Fig. 4); head tri-
angular, slightly wider (9.3) than long (9.2); snout obtusely pointed, rounded in profile, projecting beyond lower
jaw; eye small, shorter (2.3) than snout (3.2); canthus rostralis rounded, straight; lore sloping, weakly concave; nos-
tril lateral, on canthus rostralis, closer to tip of snout (1.4) than to eye (1.6); interorbital distance (3.0) wider than
internarial distance (2.5), the latter twice width of upper eyelid (1.1); pineal spot absent; tympanum hidden; upper
jaw edentate; tongue oval, without papillae; slit-like openings to a median subgular vocal sac.
Forelimb short (10.3) and stout; fingers thick, free of web, but with slight skin fringes on both sides of second
and third and inner side of fourth; second finger slightly longer (measured from inner side 2.0, outer 1.4) than
fourth (inner 1.6), latter much longer than first (outer 0.7); tips of three outer fingers weakly dilated and forming
weak disks slightly wider than basal phalanges, dorsally with median longitudinal groove; diameter of first finger
(0.3) one-third that of third finger disk (0.9), latter less than two-times width of phalange; a single outer palmar
tubercle (1.1) much larger than inner (0.6); distinct, rounded subarticular tubercles, numbering one on first and sec-
ond fingers, two on third and fourth fingers; nuptial pad absent (Fig. 5).

FIGURE 4. Dorsal (A) and ventral (C) views of male holotype of Microhyla malang sp. nov. (KUHE 17245) and dorsal view
(B) of syntopic male Microhyla borneensis (KUHE 53814) from Gunung Serapi, Sarawak. Scale bar = 10 mm.

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FIGURE 5. Ventral views of right hands (A, C) and feet (B, D) of syntopic Microhyla borneensis (A, B: KUHE 53814) and
Microhyla malang sp. nov. (C, D: KUHE 53018) from Gunung Serapi, Sarawak. Scale bar = 5 mm.

FIGURE 6. Dorsolateral view of a male paratype of Microhyla malang sp. nov. (KUHE 53018) from Gunung Serapi, Sarawak.

Hindlimb long (40.0) about three times length of forelimb; tibia long (14.0), heels overlapping when limbs are
held at right angles to body; tibiotarsal articulation of adpressed limb reaching far beyond snout tip; foot (12.4)
shorter than tibia; tips of toes distinctly dilated into disks, much wider than those of fingers (disk diameter of third
toe 0.9), dorsally with median longitudinal groove; third toe longer than fifth; webs between toes moderately devel-
oped (Fig. 4), formula: I 1–2 II 1–22/3 III 11/2 –3 IV 3–1 V; subarticular tubercles prominent, rounded, numbering
one on first and second toes, two on third and fifth toes, and three on fourth toe; inner metatarsal tubercle oval,
length (0.7) one-third of first toe (2.2); outer metatarsal tubercle elevated, larger (0.9) than inner one.
Skin above with scattered, low tubercles; eyelid without supraciliary spines; a weak, curved supratympanic
fold from eye to axilla; side of body scattered with tubercles or low ridges more densely than dorsum; hindlimb
dorsally scattered with few tubercles; ventral side of body and limbs smooth.

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 41
Color. Color in life pinkish brown dorsally, with a darker wide interorbital bar, reverse triangle in shape, medi-
ally followed by three darker markings narrowly outlined in white, consisting of two wide-limbed reverse V-
shaped marks from occiput to sacrum, and a subtriangle mark between sacrum and groin; a narrower and paler line
paralleled with each dark marking laterally to flank; a small black spot at middle of two wide-limbed marks, sur-
rounded by dark dots; lips spotted with black; a cream stripe extending from eye to axilla bordered above by
supratympanic fold and below by dark labial marks (Figs. 6, 7); a narrow, broken, black lateral stripe extending
from above arm to half length of trunk; limbs dorsally with narrow dark brown bars; median edge of tarsus darkly
marked; a dark spot surrounding anus; throat mottled with dark brown and abdomen cream white; underside of
limbs scattered with black pigmentation; iris dorsally and ventrally golden with black reticulation, darkly pig-
mented at anterior and posterior corners. In preservative, pattern has not obviously changed, although color has
slightly faded.

FIGURE 7. Dorsolateral view of head of Microhyla borneensis (A–B, males [A: KUHE 53938; B: KUHE 10692]: C–F,
females [C: holotype, BM 1947. 2.11.82; D: KUHE 53020; E: KUHE 12067; F: KUHE 53165] and Microhyla malang sp. nov.
(G–H, males [G: holotype, KUHE 17245; H: KUHE 17217]; I, female [KUHE 44228]).

Variation. Individuals of the type series are generally similar in morphology. Variation in size and body pro-
portions is given in Table 2. Web formula was stable except for the third toe, which showed slight variation; on the
inner side, some had only two phalanges free of web, while on the outer side, many had 12/3 phalanges free (Fig. 1).
Tadpoles. A total of ten tadpoles from St. 25 (TOTL = 9.0 mm, HBL = 2.9 mm) to 41 (15.9 mm, 5.1 mm) from
near the type locality of M. malang and identified as that species by DNA analyses (see above) were closely exam-
ined (Table 3). Head and body flattened above, spheroidal below; HBW maximum at level of eye 58–64% (median
= 61%) of HBL; HBD 84–100% (median = 89%) of HBW; snout broadly rounded, almost truncate in profile; eyes
lateral, visible from below, eyeball 14–17% (median = 15%) of HBL; interorbital wide, 180–256% (median =
223%) of eyeball diameter; eye-snout distance 27–30% (median = 30%) of HBL. Oral disk dorso-terminal; lower
lip moderately expanded with width 33–48% (median = 43%) of HBW, with a prominent lateral papilla (sensu Das
and Haas, 2010) on each side; labial teeth and jaw sheaths entirely absent, but lower labium scattered with small

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papillae on lateral margin. Spiracle opening median, without free flap, opening 79–90% (median = 84%) of dis-
tance from tip of snout to end of body; vent median, in form of long tube directed obliquely backward, small open-
ing at edge of ventral fin; gut in form of two, thick loops visible ventrally. Tail long and lanceolate, abruptly
tapering in posterior half and drawn out into a long filament in posterior one-fourth; tail length 176–211% (median
= 205%) of HBL, maximum depth 19–28% (median = 25%) of length; dorsal fin originating at end of head-body,
with a straight margin, sub-parallel with much deeper ventral fin in anterior half of tail; ventral fin deeper than dor-
sal throughout anterior to tail filament; caudal muscle moderately strong, maximum tail width 28–36% (median =
32%) of HBW; muscle depth maximum at origin, 41–55% (median = 48%) of maximum tail depth, but steadily
narrowed posteriorly, with depth at middle of tail shallower than upper fin depth. Color in life (Fig. 8) light brown
on dorsum and laterally, with a pair of paler interorbital markings and darker bands on middle and end of body;
venter grey and belly translucent in young stages; tail at middle with two black markings separated by an oblique
white marking; tail tip transparent.

TABLE 2. Measurements in 12 males of Microhyla malang sp. nov. SVL (Mean+1SD, in mm) and medians of ratios (R) of other
characters to SVL, followed by ranges in parenthesis. See text for character abbreviations.

SVL RHL RSL REL RN-EL RHW


Mean/Median 20.5+1.2 39.5 15.8 12.1 7.0 41.0
Range (18.7–22.2) (37.9–42.9) 䋨14.4–17.6) 䋨10.9–12.8) (6.2–8.0) (38.3–43.6)
RIND RIOD RUEW RFLL RLAL RHAL
Median 11.4 13.5 6.2 52.2 38.4 22.8
Range (10.5–11.7) (12.3–15.3) (5.2–6.9) (48.2–54.1) (37.1–42.0) (22.2–23.7)
R1FL RIPTL ROPTL RHLL RTL RFL
Median 5.4 2.8 4.3 193.8 65.9 58.2
Range (5.2–6.3) (2.2–3.2) (3.6–5.5) (182.1–199.7) (62.3–68.8) (53.2–61.0)
R1TOEL RIMTL ROMTL R1FW R2FDW R3FDW
Median 10.4 4.0 5.8 1.6 3.0 3.6
Range (9.2–11.7) (3.1–4.5) (4.2–7.0) (1.1–2.8) (2.3–3.4) (2.7–4.5)
R4FDW R1TDW R2TDW R3TDW R4TDW R5TDW
Median 3.1 3.7 5.6 6.2 5.8 4.8
Range (2.7–3.6) (2.9–4.7) (4.2–6.3) (4.6–6.7) (4.6–6.7) (4.1–5.7)
R1FLO R2FLI R2FLO R3FLI R3FLO R4FLI
Median 2.9 9.6 6.6 13.6 13.2 7.3
Range (2.4–4.1) (7.7–10.6) (5.8–7.1) (12.7–14.9) (11.7–13.8) (5.6–8.2)

Range. The new species is so far known from western Sarawak and eastern Sabah, Malaysian Borneo. Sara-
wak: Gunung Serapi, Matang, (the type locality), Kanowit, and Lanjak Entimau; Sabah: Poring Hot Spring, Sepi-
lok (Matsui 1979 as M. borneensis), Tabin, and Tawau. The species ranges in altitude from 50 to 555 m a.s.l. Das &
Haas (2010) reported Gunung Poing, Bau (Sarawak), Batu Apoi Forest Reserve, Temburong (Brunei Darussalam),
and Kalimantan (Indonesia) as the range of their M. borneensis (= M. malang).
Natural history. As already reported by Das and Haas (2010), M. malang is sympatric with M. borneensis
around the type locality, in Kubah National Park. They are actually syntopic and can be found very close to each
other near the Frog Pond of the park. Chorus of male M. malang was heard in August, September, and December in
the park, and also reported from there in April (Dehling 2010). Calls were heard also in August in Kanowit and
Tawau. Tadpoles of two different stages were found in August in Kubah, and older ones in Sepilok, Tabin, and
Tawau in August. Matsui (1979) reported a female (now KUHE 44228) from Poring, Sabah with enlarged ova in
March. Thus M. malang seems to have a prolonged breeding season. Tadpoles were found in ponds and in shallow,
muddy pools in drying stream beds.
Call characteristics. Characteristics of calls recorded at the type locality did not differ from those already
reported by Dehling (2010) from records of the same locality at air temperatures of 23.5−24.7°C. They consisted of

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 43
notes each emitted at an interval of ca. 830 ms. Each note was composed of 5.7 ± 1.1 (4−8) short pulses and lasted
for 169 ± 37 (104−242) ms. Frequency bands spread over the 1400–2900 hz range, and the dominant one was 2404
± 94 (2250−2530) hz. Frequency modulation was not marked but weak intensity modulation was present (Dehling
2010).

TABLE 3. Measurements in mm of larval Microhyla malang sp. nov. from Serapi. See text for character abbreviations.
Stage 25 27 30 31 31 32 34 35 40 41
TotL 9.02 12.90 13.11 13.66 14.42 13.86 13.67 14.90 15.94 15.85
HBL 2.93 4.15 4.28 4.78 5.11 4.89 4.96 4.88 5.14 5.09
HBW 1.86 2.48 2.75 2.85 3.12 3.03 2.88 3.11 3.09 3.14
HBD 1.66 2.31 2.31 2.47 2.88 2.80 2.88 2.75 2.69 2.75
Eye-Snout 0.90 1.27 1.22 1.35 1.50 1.47 1.36 1.51 1.49 1.54
Eye Diameter 0.50 0.58 0.68 0.71 0.72 0.69 0.69 0.77 0.80 0.79
Inter-orbital 0.90 1.21 1.54 1.56 1.73 1.77 1.67 1.76 1.60 1.65
Snout-Spiracle 2.48 3.68 3.52 4.30 4.15 4.00 3.92 4.14 4.34 4.29
Oral disk W 0.66 1.04 1.15 1.29 1.56 1.35 1.38 1.27 1.02 1.38
TailL 6.09 8.75 8.83 8.88 9.31 8.97 8.71 10.02 10.80 10.76
Max tail D 1.51 2.04 2.20 2.15 2.65 2.34 2.48 2.42 2.06 2.31
Max tail W 0.62 0.89 0.79 0.86 1.04 1.04 0.87 1.05 0.86 0.94
Max muscle D 0.83 0.92 1.10 1.02 1.15 1.09 1.04 1.16 1.14 1.21

Comparisons. As noted above, the new species closely resembles, but can be differentiated from M. borneen-
sis by larger body size (male SVL 19.4–21.3 mm vs.11.0–13.2 mm in M. borneensis), more developed toe web
(fifth toe with one phalange free of web vs. two phalanges free), possession of (vs. usually lacking) a mid-dorsal
black spot and light lines at the edge of dorsal dark markings. They are remarkably different in tadpole morphology
and ecology. The new species breeds in ponds and pools of slowly flowing streams, while M. borneensis utilizes
small amount of the water in the cups of pitcher plants. Tadpoles of the new species reach larger size (15.9 mm in
total length vs. 11.5 mm in M. borneensis), and have darker body color with dorsal and tail markings (nearly trans-
parent and without marking in M. borneensis). The new species is morphologically similar to M. mantheyi, with
which it has long been confused (as M. borneensis). However, M. malang is distinguishable from M. manthey by its
uniformly brown snout and forehead (vs. top of snout cream colored in M. mantheyi), shorter dark, lateral stripe
covering anterior half of flank (vs. longer stripe covering entire flank), pale supratympanic stripe lacking dark mar-
gin (vs. pale stripe with a dark dorsal edge), absence of light vertebral line (vs. presence of a thin line), rounded
snout (vs. snout tapering), and fourth toe three phalanges free of web (vs. fourth toe more than three phalanges
free). In tadpoles, M. malang with a light interorbital bar and without dark terminal filament of tail also differs from
M. mantheyi, in which interorbital bar and tail filament are dark. Although Das & Haas (2010: 45) noted absence in
M. mantheyi of finger disks, the species has disks like those of M. malang and M. borneensis as clearly stated in the
original description (Das et al. 2007a). The new species differs from all the other congeners in the same manner as
was reported for M. borneensis by Das & Haas (2010, as M. nepenthicola).
Calls of M. malang and M. borneensis are basically similar in temporal structure, but clearly differ in fre-
quency properties with the new species emitting a lower voice than M. borneensis. Calls of sympatric M. borneen-
sis recorded on 22 January 1990 at an air temperature 21.7 C included two types. The first call type consisted of a
series of pulsed notes lasting 50–140 ms and containing 2–4 pulses. The second call type consisted of the first call
type and additional notes including longer series of pulsed notes, and lasted 420–510 ms. The additional note was
composed of 6–8 pulses. The pulse rate in the first call type ranged from 39.2–41.6 per second. The dominant fre-
quency was spread over the 3460–3750 hz range. A well-defined harmonic was seen at about 4625 hz and fre-
quency or intensity modulations were absent.

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FIGURE 8. Larval Microhyla malang sp. nov. from Gunung Serapi, Sarawak (A–C: stage 32, total length = 13.9 mm) and
Tawau, Sabah (D–F: stage 41, total length = 16.4 mm, from photos taken by T. Shimada); dorsal (A, D), lateral (B, E), and ven-
tral views (C, F). Scale bar = 5 mm.

Discussion

It is clear that Das & Haas (2010) erroneously considered true M. borneensis as a new species (= M. nepenthicola),
without noticing the unnamed status of their M. borneensis (= M. malang). All illustration guides currently avail-
able show the picture and body size of M. malang as those of M. borneensis (e.g. Inger & Stuebing 1997; Malkmus
et al. 2002; Das 2007), most probably following the description of Inger (1966). However, confusion of M. malang
with M. borneensis seems to date back to Parker's action.
The original description of M. borneensis (Parker 1928: 481) was somewhat modified in the later microhylid
monograph (Parker 1934: 128; see Appendix 2). Of the many changes made, the most important ones include
degree of toe web development, state of dorsal dark marking, and body size. Toe web development in the original
description, "between 1/2 and 2/3 webbed", was changed to "rather more than half webbed", and the original notation
that "the web leaving two phalanges of the third and fifth toes" was deleted. Next, for dorsal dark marking ("ill-
defined darker area" or "spot"), addition was made that "it may be outlined in white". Also, addition of "a light line
from behind eye to the forelimb" was made in the monograph. However, the most significant change is the SVL,
from the original 18 to 23 mm.
Related to these changes is the addition of specimens examined in the monograph. A female from Mt. Dulit,
Sarawak (BM 1933.6.20.15), as well as a developmental series of larvae (BM 1914.5.12.61–78), were added to a
single female holotype (unfortunately, I could not locate them during my visit). From this fact, it is considered that
the changes in the monograph would have been made on the basis of the newly obtained female which should actu-
ally be not conspecific with M. borneensis. Larger body size (SVL = 23 mm) and white-edged dorsal dark marking
and light postoccular marking surely represent diagnostic characteristics of M. malang, sp. nov. Deletion of the
phrase, "web leaving two phalanges of the third and fifth toes" in the monograph is interpreted to indicate better
developed toe webbing of the additional specimen. Most probably, Parker (1934) erroneously confused these two
very similar species, and modified his description of M. borneensis in the monograph. This last publication alone
seems to have been referenced by the subsequent researchers, and led to the currently widespread misidentification
of M. malang.

TAXONOMIC REVISION OF ONE OF OLD WORLD’S SMALLEST FROGS Zootaxa 2814 © 2011 Magnolia Press · 45
Although Das et al. (2007a) split M. mantheyi from M. borneensis (actually M. malang), Das & Haas (2010)
did not include this Peninsular species in their phylogenetic analyses. Our analyses indicated nearly monophyletic
relationships of the three species, within which two Bornean species are sister to each other. This indicates that a
common ancestor of M. malang and M. borneensis began speciation in Borneo after it was split from ancestral M.
mantheyi that occurred in the present Malay Peninsula. Because M. malang and M. mantheyi are open water breed-
ers as typically found in many other anuran species, while M. borneensis is a unique phytotelmic breeder, ecologi-
cal specialization of M. borneensis within Borneo is also strongly suggested. Microhyla malang proved to be
somewhat differentiated genetically among local populations, and might have a long history of differentiation on
the island. Unfortunately, no comparable data are available for M. borneensis. As noted above M. malang can be
sympatric or even syntopic with M. borneensis around the type locality in Sarawak. However, in other localities
(Kanowit and Lanjak Entimau in Sarawak, and Kinabalu, Sepilok, Tabin, and Tawau in Sabah), occurrence of only
M. malang has been confirmed. It might be thought that the unique breeding habits of the species might be restrict-
ing range expansion within Borneo. However, the pitcher plant Nepenthes ampularia, in which the species usually
breeds, occurs widely in the island from lowland to high elevations, including disturbed areas. The small size of the
frog has most likely prevented it from being noticed by researchers and its occurrence might eventually be revealed
in wider regions of Borneo.
Microhyla borneensis was originally allied to the M. berdmorei group by Parker (1928, by implication), and
the species (including M. malang) has been placed in the same group of the subgenus Microhyla (Dubois 1987).
Our results from a limited number of species did not support such a grouping. A clade including three species
related to M. borneensis showed unresolved relationships with M. berdmorei, as well as M. okinavensis of the
Dubois’ (1987) subgenus Diplopelma. Likewise, M. achatina was not monophyletic with M. heymonsi, placed in
the M. achatina group by Dubois (1987), but with M. malang, M. borneensis, and M. mantheyi. Thus, Parker's
(1928) and Dubois’ (1987) groupings of the genus Microhyla based on morphology require thorough reexamina-
tion.
Das & Haas (2010) characterized, M. nepenthicola (= true M. borneensis) as "Old World's smallest frog". This
statement requires some comments and modifications. Actually another Bornean congener, M. perparva (10.1–
10.5 mm in males and 11.4–12.4 mm in females: my measurements), is smaller than M. borneensis when females
are taken into consideration, and the female of M. superciliaris from Peninsular Malaysia and Sumatra are also
smaller (11.8 mm, my measurement of the holotype). Other, similarly miniaturized frogs of the genera Stumpffia
and Sooglossus also occur in the Old World (Madagascar and Seychelles).
In relation to miniaturization, Das and Haas (2010) associated reduction in the first finger of M. borneensis
with navigation on the slippery zone of Nepenthes pitchers. This idea presents some difficulties. First, reduction of
the first finger is more pronounced in males than in females. Because females in this species are much larger than
males, it would be more difficult for them to navigate in the pitcher than males. Moreover, amplexus is made out-
side of the pitcher, and, as is the case in most anurans, the movement of the amplectant pair is promoted only by the
female (my own observation). Therefore, females are expected to require a higher degree of reduction in their fin-
gers than males, which is actually not the case. Second, in other diminutive Bornean congeners, M. perparva is
reported to breed in rain pools in the forest (Inger & Stuebing 1997), and M. petrigena, with male SVL of 14–16
mm and also categorized as diminutive according to Das & Haas (2010), breeds in small potholes on rocky banks
of streams (Inger & Stuebing 1997; my own observation). In these two species, reduction of the first finger is more
distinct than in M. borneensis, and they are even noted to have just three fingers (Inger & Stuebing 1997). Thus,
reduction in the first finger of these diminutive species might be not directly related to their breeding habits, and
the hypothesis of Das and Haas (2010) requires reconsideration.
The results of my own observation of larval M. borneensis, such as small total length and immaculate body and
tail, conform to descriptions by both Parker (1934) and Das & Haas (2010, as M. nepenthicola). Although there are
some discrepancies such as his mention of narial openings in Parker's (1934) description, this author can be thought
to have treated the same species of larvae as did Das & Haas (2010). Thus, identification by Parker (1934) of
nepenthiphilous larvae as M. borneensis is justified. Oddly, Das & Haas (2010: 47) rejected Parker' s (1934) identi-
fication simply on the basis of their historical review of the origin of specimens, and not from comparison of
descriptions. Description of larval M. borneensis by Inger (1966) seems to have been followed by Malkumus et al.
(2002), but because adult and larval association of M. borneensis and nepenthiphilous larvae is established now,
Inger's (1966) larvae could be considered as that of M. malang. However, they are different in total length (26.1–

46 · Zootaxa 2814 © 2011 Magnolia Press MATSUI


32.0 mm in Inger's larvae vs. <15.9 mm in larval M. malang), and this difference is too large to be ascribed to vari-
ation caused by different environmental conditions. Our limited samples from Tawau and Tabin, Sabah, are simi-
larly small, suggesting constant small size in larval M. malang. In addition to this, many characters such as type of
spiracle, position of spiracle opening, condition of vent , and tail coloration, differ markedly between Inger’s
(1966) larvae and larval M. malang. It would be therefore safe to consider Inger's tadpole to be that of another spe-
cies of Microhyla.
What is interesting is that available references denote that funnel-shaped mouth is seen only in M. achatina, M.
heymonsi, and M. mixtura (e.g. Parker 1928, 1934; Inger 1966; Fei et al. 2005). In examining larval Microhyla, I
found the oral morphology of M. malang shows an intermediate condition between M. heymonsi and other species
including M. berdmorei and M. petigena. Unfortunately, I could not examine larval M. achatina, but description of
oral apparatus for that species seems to be partially applied to larval M. malang. Future detailed examination of
oral apparatus in this genus surely will give some insights into many problems, including niche segregation among
congeners.

Acknowledgements

The Economic Planning Unit, Prime Minister’s Department (former Socio-Economic Research Unit) of Malaysia,
the State Government of Sarawak, the Forest Department, Sarawak, the Sarawak Forestry Corporation, and Sabah
Parks kindly permitted me to conduct the project, and Universiti Malaysia Sabah, the Japan International Coopera-
tion Agency, and the Forest Department, Sarawak kindly provided all the facilities for conducting research. I am
grateful to the following for their encouragements and/or permission to conduct research and /or field companion-
ship: the late A.H. Abang, K. Araya, L. Chong, Datuk C. Ek Choon, B. Daicus and his family, K. Eto, T. Hayashi,
the late T. Hidaka, T. Hikida, A.H. Kassim, J.J. Kendawong, the late T. Kobayashi, K.B. Kueh, T. Kusano, D.
Labang, M.B. Lakim, R. Mohidin, A. Mori, J. Nais, K. Nishikawa, C.-K. Onn, Datuk Haji Len Talif Salleh, L.-H.
Seng, T. Shimada, T. Tachi, N.-S. Wong, and Haji ali Bin Yusop. I am also indebted to A. Hamidy and N. Kuraishi
for laboratory assistance, and to C. McCarthy and B. T. Clarke (BM), A. Dubois and A. Ohler (MNHNP), A.
Norhayati (UKM), M. Maryati and A. Sudin (UMS), P. Yambun (SP), and R. Zainudin (UNIMAS) for allowing me
to examine specimens under their care. Field trips were made possible by grants from The Monbusho International
Scientific Research Programs and from The Monbusho through the Japan Society for the Promotion of Science
(Field Research, Nos. 01041051, 02041051, 03041044, 04041068, 10041166, 15370038, 20405013). I thank T.
Johnson for reviewing English text.

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48 · Zootaxa 2814 © 2011 Magnolia Press MATSUI


APPENDIX 1.

Specimens used for morphological comparisons. BM = Natural History Museum, London; KUHE = Graduate School of
Human and Environmental Studies, Kyoto University; MNHNP = Museum National d'Histoire Naturelle, Paris; RMNH =
Rijksmuseum van Natuurlijke Historie, Leiden; SM = Sarawak Museum; ZMB = Zoologisches Museum, Universitåt
Humboldt, Berlin.

Microhyla achatina: BM 1947.2.11.59 (cotype); RMNH 1728 (5 syntypes)


Microhyla annamensis: BM 1947.2.11.60 (holotype), 61–63 (paratypes)
Microhyla annectens: KUHE 52438, 53372–74
Microhyla berdmorei (including Microhyla fowleri): KUHE 52034–35, 52373, 52386–95, 53363–67
Microhyla borneensis: BM 1947. 2.11.82; SM unnumbered (former KUHE 53167, 53814); KUHE 10689–95, 10742–44,
12039–40, 12067, 12070, 26417–19, 53020, 53033, 53165, 53241, 53840, 53852, 53938
Microhyla butleri: BM 1947.2.11.70 (holotype), KUHE 19636–38, 19652–56, 19956–58, 52033, 52560
Microhyla fissipes: BM 1947.2.11.85 (holotype); KUHE 23065–72, 23767–69, 23887–88, 52431, 53256–59, 53273, 53304,
53487-48
Microhyla heymonsi: ZMB 219441; KUHE 53256–53259, 53360–62
Microhyla mantheyi: KUHE 15726–27, 35440, 53367–69, 53487, 53492
Microhyla marmorata: KUHE 32455
Microhyla okinavensis: KUHE 50016, 50023, 50029, 50418–20, 50431–32, 50439-40
Microhyla ornata: MNHNP 2418.5 (holotype)
Microhyla palmipes: BM 1947.2.11.73 (holotype)
Microhyla perparva: KUHE 10399–400, 10454, 10459, 53625
Microhyla petrigena: KUHE 53743–47
Microhyla pulchra: KUHE 19762, 19765, 19774, 19776, 19793
Microhyla superciliaris: BM 1947.2.11.81 (holotype)

APPENDIX 2

Original description (Parker 1928) and changes in the microhylid mognograph (Parker 1934) of Microhyla borneensis. Under-
lined words in the original description were deleted and changed to or added by the italicized words in parentheses in the
monograph.

Snout rounded (pointed), slightly projecting (prominent), once and a quarter (half) as long as the diameter of the eye. (canthus
rostralis rounded; loreal region slightly oblique; interorbital space once and three-quarters to twice and a half as broad as
the upper eyelid; tympanum hidden). Fingers and toes moderate, dilated distally into (with) distinct disks with a median
groove on their upper surface; first finger very short; fourth longer than second (second shorter than the fourth; toes with
disks similar to, but larger than, those of the fingers;) toes between 1/2 and 2/3 (rather more than half) webbed, the web
almost reaching the discs of the first and second toes on their inner margin, but leaving two phalanges of the third and fifth
toes and three of the fourth toe free (membrane midway between the third and fourth, extending half-way between the
tubercles of the third toe; fifth toe a little shorter than the third); subarticular tubercles well developed; two metatarsal
tubercles, the (an oval) inner elongate, the (and a conical outer conical (metatarsal tubercle). Tibio-tarsal articulation
reaching somewhat (a little) beyond the end (tip) of the snout; tibia 3/5 the length of the head and body. Skin smooth
(above and below); a slight (tympanic) fold from the posterior angle of the eye to the fore limb. Above light brown with an
ill-defined darker area along the middle of the back; this stretches from (the head, where it connects) the upper eyelid<s>
to eyelid, being truncate anteriorly, (it) narrows behind the occiput, broadens and gives off a short branch towards the fore
limbs (or the flanks), narrows (again) on the middle of the back, and (finally) widens and becomes indistinct behind this
point (on the posterior part of the back; this spot may be outlined in white); A dark brown (, oblique) streak from above the
shoulder to the middle of the flank. Hind limbs very indistinctly cross-banded (barred); a dark spot on the front of the
knee. A dark, light-edged area above (about) the vent. Upper lip with three dark brown spots (and a light line from behind
eye to the fore-limb). Lower surfaces yellowish white, faintly mottled with light brown. Snout to vent (length) 18 (23) mm.
Fore limb 8 mm. Hind limb 31 mm.

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