A new species of the genus Pulchrana Dubois, 1992 (Amphibia, Ranidae) from Sumatra, Indonesia

Abstract

We describe a new species of Pulchrana from the island of Sumatra, in western Indonesia. Pulchrana fantastica, new species, is currently only known from Aceh and Sumatera Utara provinces. It is related to P. siberu and P. centropeninsularis as inferred from morphological similarity and phylogenetic relationships, estimated from DNA sequences. It can be diagnosed by the following unique combination of characters: (1) SVL adult males 40.32–45.19 mm; (2) males have large humeral glands on the anteroventral surface of brachium, nuptial pads absent; (3) dorsal skin finely granular to granulated, with or without asperities at the tip of granules; (4) webbing formula:
I(1 ― 1–11/2)II(1/2 ― 2)III(1 ― 2–21/3)IV(2–21/3 ― 1)V; (5) straight dorsolateral stripes, thin, continuous, anteriorly confluent at snout, posteriorly interconnected by a series of spots; (6) middorsum in adults black with light medial line or combination of spots and line, black without marking in juveniles; (7) dense cream or yellow to orange spots on flanks, and dorsal side of limbs, spots non-uniform in shape; (8) venter greyish or brown, with small light dots on throat and chest, sometimes reaching abdomen; (9) iris background black, lower part with dense orange stippling, upper region with orange reticulation with gold in the middle, pupil encircled with solid orangegolden line; (10) upper and lower lip grey or brown with cream or yellow spots. We also report the presence of P. centropeninsularis from Sumatra; a species previously known only from a single locality in the Malay Peninsula; and discuss the biogeographic implications of this significant range extension.

Full text

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RAFFLES BULLETIN OF ZOOLOGY 2018
INTRODUCTION
Despite being renowned for its high herpetofaunal diversity,
the island of Sumatra remains severely understudied (e.g.,
Iskandar & Colijn, 2000; Stuart et al., 2006; Inger et al., 2009).
Many new species of anuran amphibians have been described
from Sumatra in the last decade, such as Duttaphrynus totol
(Teynie et al., 2010), Sigalegalephrynus mandailinguensis,
S. minangkabauensis (Smart et al., 2017), Limnonectes
sisikdagu (McLeod et al., 2011), Pulchrana rawa (Matsui et
al., 2012), Sumaterana dabulescens, S. montana (Arin et al.,
2018), Chiromantis nauli, C. baladika (Riyanto & Kurniati,
2014), Rhacophorus indonesiensis (Hamidy & Kurniati,
2015), Philautus amabilis, P. polymorphus, P. thamyridion,
and P. ventrimaculatus (Wostl et al., 2017). Nevertheless,
a substantial portion of Sumatra’s anuran diversity likely
remains hidden within common, morphologically cryptic, and
widespread species (Stuart et al., 2006; Inger et al., 2009).
A new species of the genus Pulchrana Dubois, 1992 (Amphibia: Ranidae)
from Sumatra, Indonesia
Umilaela Arin1*, Ganjar Cahyadi2, Utpal Smart3, André Jankowski1, Alexander Haas1
Abstract. We describe a new species of Pulchrana from the island of Sumatra, in western Indonesia. Pulchrana
fantastica, new species, is currently only known from Aceh and Sumatera Utara provinces. It is related to P. siberu
and P. centropeninsularis as inferred from morphological similarity and phylogenetic relationships, estimated from
DNA sequences. It can be diagnosed by the following unique combination of characters: (1) SVL adult males
40.32–45.19 mm; (2) males have large humeral glands on the anteroventral surface of brachium, nuptial pads absent;
(3) dorsal skin nely granular to granulated, with or without asperities at the tip of granules; (4) webbing formula:
I(1 ― 1–11/2)II(1/2 ― 2)III(1 ― 2–21/3)IV(2–21/3 ― 1)V; (5) straight dorsolateral stripes, thin, continuous,
anteriorly conuent at snout, posteriorly interconnected by a series of spots; (6) middorsum in adults black with
light medial line or combination of spots and line, black without marking in juveniles; (7) dense cream or yellow
to orange spots on anks, and dorsal side of limbs, spots non-uniform in shape; (8) venter greyish or brown, with
small light dots on throat and chest, sometimes reaching abdomen; (9) iris background black, lower part with dense
orange stippling, upper region with orange reticulation with gold in the middle, pupil encircled with solid orange-
golden line; (10) upper and lower lip grey or brown with cream or yellow spots. We also report the presence of P.
centropeninsularis from Sumatra; a species previously known only from a single locality in the Malay Peninsula;
and discuss the biogeographic implications of this signicant range extension.
Key words. molecular systematics, morphology, new species, Pulchrana signata Complex, Sumatra, taxonomy
RAFFLES BULLETIN OF ZOOLOGY 66: 277–299
Date of publication: 3 May 2018
http://zoobank.org/urn:lsid:zoobank.org:pub:807DCF10-9529-4C53-9190-74E5004CDDDB
© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)
1Centrum für Naturkunde-Zoologisches Museum Hamburg, Universität Hamburg,
Edmund-Siemers-Allee 1, 20146 Hamburg, Germany; Email: umilaela@gmail.com,
umilaela.arin@studium.uni-hamburg.de (*corresponding author), andre.jankowski@
zoologie.uni-hamburg.de, alexander.haas@uni-hamburg.de
2School of Life Sciences and Technology, Institute of Technology Bandung, Jalan
Ganeca 10 Tamansari Bandung 40132 Indonesia; Email: ganjar@sith.itb.ac.id
3Amphibian & Reptile Diversity Research Center, Department of Biology, University
of Texas at Arlington, TX 76019 USA; Email: utpal.smart@uta.edu
The Pulchrana signata group is one such complex of
relatively abundant ranid frogs, distributed across Sundaland
and the Philippines with at least nine nominal species (Brown
& Guttman, 2002; Brown & Siler, 2013; Chan et al., 2014;
Oliver et al., 2015). In total, the genus Pulchrana currently
includes 16 species that were formerly subsumed under the
genus Hylarana (Frost, 2018). The most recently described
species are P. centropeninsularis from the Malay Peninsula
(Chan et al., 2014) and P. guttmani from Mindanao Island
of the southern Philippines (Brown, 2015). The rst of these
taxa initially had been assigned to the name P. siberu (Leong
& Lim, 2004), based on phenotypic similarity to an endemic
taxon from the Mentawai Islands (Dring et al., 1990). The
addition of genetic data (Brown & Siler, 2013) revealed this
to be a distinct species based on phylogenetic relationships.
In a subsequent analysis, Chan et al. (2014) reported the
presence of another hitherto undescribed lineage from the
Batak Mountains of Sumatera Utara Province, which was
closely related to, yet also markedly distinct from both P.
siberu and P. centropeninsularis.
During eldwork on the island of Sumatra between the
years 2013–2014, several individuals belonging to the genus
Pulchrana of uncertain taxonomic afnities, were collected
from Aceh and Jambi provinces. Upon further investigations,
based on molecular and morphological approaches, these
specimens were identied as two taxa: P. centropeninsularis
and the hitherto unnamed species mentioned in Chan et al.
(2014). In this paper we provide a formal species description
for the latter, and we justify the recognition of the new species
Taxonomy & Systematics

278
Arin et al.: New Pulchrana from Sumatra
Table 1. List of morphometric characters, acronyms, character denitions (and their citations) used in this study.
Primary Name Primary Acronym Denition Citation
Anterior Eye to Eye Distance EED The distance from anterior of left eye to the
anterior of right eye
Chan et al., 2014
Brachium Length BL The distance from axilla to exed elbow Chan et al., 2014
Disc Width of Finger I, Finger
II, Finger III, Finger IV
Fin1DW, Fin2DW,
Fin3DW, Fin4DW
The widest horizontal diameter of disc of
Finger I, Finger II, Finger III, Finger IV
Watters et al., 2016
Disc Width of Toe I, Toe II,
Toe III, Toe IV, Toe V
Toe1DW, Toe2DW,
Toe3DW, Toe4DW,
Toe5DW
The widest horizontal diameter of disc of
Toe I, Toe II, Toe III, Toe IV, Toe V
Modied from Watters
et al., 2016
Dorsolateral Stripe Width DLSW Maximum width of dorsolateral stripe,
recorded above tympanum
Brown & Guttman,
2002
Eye Diameter ED The distance between anterior and posterior
corner of upper and lower eyelids
Chan et al., 2014
Eye-Nostril Distance END The distance from anterior of eye to the
posterior of nostril
Chan et al., 2014
Femur Length FL The distance from vent to outer margin of
exed knee
Chan et al., 2014
Finger I Length Fin1L The distance from proximal margin of the
inner metacarpal to the tip of the Finger I
Duellman & Trueb,
2015
Finger II, Finger III, Finger IV
Length
Fin2L, Fin3L, Fin4L The distance from proximal margin of the
palmar tubercle to the tip of the Finger II,
Finger III, Finger IV
Watters et al., 2016
Forearm Length FAL The distance from exed elbow to base of
inner metacarpal tubercle
Chan et al., 2014
Head Length HL The distance from posterior margin of lower
jaw to tip of snout
Chan et al., 2014
Head Width HW The distance taken immediately from
posterior to eyes
Chan et al., 2014
Humeral Gland Length HG The horizontal length of humeral gland Chan et al., 2014
Internarial Distance IND The shortest distance between the inner
margins of the nostrils
Chan et al., 2014
Interorbital Distance IOD The distance across top of head between
medial margins of orbits at their closest
points
Chan et al., 2014
Inner Metatarsal Tubercle
Length
IMTL Greatest length of inner metatarsal tubercle Modied from Watters
et al., 2016
Inner Metatarsal Tubercle
Width
IMTW Greatest width of inner metatarsal tubercle Modied from Watters
et al., 2016
Inner Metacarpal Tubercle
Length
IMCL Greatest length of inner metacarpal tubercle Modied from Watters
et al., 2016
Inner Metacarpal Tubercle
Width
IMCW Greatest width of inner metacarpal tubercle Modied from Watters
et al., 2016
Length of Toe I, Toe II, Toe
III
Toe1L, Toe2L, Toe3L The distance from the base of inner
metatarsal to the tip of the Toe I, Toe II,
Toe III
Watters et al., 2016
Length of Toe IV, Toe V Toe4L, Toe5L The distance from the base of outer
metatarsal to the tip of the Toe IV, Toe V
Watters et al., 2016
Nostril-Snout Distance NSD The distance from anterior of nostril to the
tip of the snout
Chan et al., 2014

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RAFFLES BULLETIN OF ZOOLOGY 2018
Primary Name Primary Acronym Denition Citation
Outer Metatarsal Tubercle
Length
OMTL Greatest length of outer metatarsal tubercle Modied from Watters
et al., 2016
Outer Metatarsal Tubercle
Width
OMTW Greatest width of outer metatarsal tubercle Modied from Watters
et al., 2016
Outer Metacarpal Tubercle
Length
OMCL Greatest length of outer metacarpal tubercle Modied from Watters
et al., 2016
Outer Metacarpal Tubercle
Width
OMCW Greatest width of outer metacarpal tubercle Modied from Watters
et al., 2016
Palmar Tubercle Length PTL Greatest length of palmar tubercle Modied from Watters
et al., 2016
Palmar Tubercle Width PTW Greatest width of palmar tubercle Modied from Watters
et al., 2016
Snout Length SL The distance from anterior corner of the eye
to tip of snout
Chan et al., 2014
Snout-Vent Length SVL The distance from the tip of snout to vent Chan et al., 2014
Tarsal Length TL The distance from outer margin of exed
tarsus to base of inner metatarsal tubercle
Chan et al., 2014
Tympanum Diameter TD The horizontal width of tympanum as its
widest points
Chan et al., 2014
Tympanum-Eye Distance TED The distance from anterior tympanum to
posterior eye
Chan et al., 2014
Tibia Length TBL The distance from outer margin of exed
knee to outer margin of exed tarsus
Chan et al., 2014
Width of Terminal Phalange
Finger I, Finger II, Finger III,
Finger IV
Fin1TPW, Fin2TPW,
Fin3TPW, Fin4TPW
Measure at midpoint of terminal phalange
of the Finger I, Finger II, Finger III, Finger
IV
Modied from Watters
et al., 2016
Width of Terminal Phalange
Toe I, Toe II, Toe III, Toe IV,
Toe V
Toe1TPW, Toe2TPW,
Toe3TPW, Toe4TPW,
Toe5TPW
Measure at midpoint of terminal phalange
of the Toe I, Toe II, Toe III, Toe IV, Toe V
Modied from Watters
et al., 2016
based on genetic and morphological evidence. Furthermore,
our collected material of P. centropeninsularis is the rst
Sumatran record for this species and signicantly extends
its known range (formerly known only from the Malay
Peninsula).
MATERIALS AND METHODS
Morphology. We examined a total of 21 frogs from three
provinces in Sumatra (Aceh: n males = 10, n juveniles =
6; Sumatera Utara: n males = 2; and Jambi: n males =3).
No female was collected during the trip. The specimens
were xed in 4% neutral-buffered formalin and later stored
in 70% ethanol. All material examined in this study are
deposited at the Museum Zoologicum Bogoriense (MZB),
Indonesia. In the future, some of the paratypes and reference
specimens shall be deposited at the Zoologisches Museum
Hamburg (ZMH), Germany. Morphometric measurements
were only taken from adult specimens (n = 15) in order to
avoid bias due to ontogenetic variation. Sexual maturity was
determined in males by the presence of humeral glands and
vocal sacs. We followed Duellman & Trueb (1986), Brown
& Guttman (2002), and Kok & Kalamandeen (2008) for
qualitative morphology assessment (e.g., head shape, skin
texture, and colouration). For the webbing formula of toes
we adopted the approach from Savage & Heyer (1997) with
the renements suggested by Guayasamin et al. (2006).
We applied the same characters and terminology used by
Chan et al. (2014) to make measurements comparable.
Additional measurements follow Duellman & Trueb (2015)
and Watters et al. (2016), for more detailed morphological
descriptions. All measurements, acronyms, denitions, and
citation are presented in Table 1, and were taken with digital
calipers (0.02 mm precision reading). We applied standard
descriptive statistics (mean, standard deviation, range, in
mm) to summarise morphological data.
Phylogenetic analyses of molecular data. Tissue samples
of two specimens from Aceh (MZB.Amph.28891 and
MZB.Amph.28946) and two specimens from Jambi
(MZB.Amph.28765–66) were selected for molecular
work. Additionally, we included five samples from the
collection of the Museum of Vertebrate Zoology (MVZ),
University of California, Berkeley, USA from the Mentawai

280
Arin et al.: New Pulchrana from Sumatra
Islands (Siberut, Sipora, and Pagai Selatan: MVZ272082,
MVZ272086, MVZ272089–91). DNA was extracted from
liver samples (preserved in either 96% ethanol or RNAlater)
using DNA Analytic Jena® Kit (Germany). We made 20
µL PCR reactions as follows: 10 µL Green Taq Promega,
8 µL H2O, 0.5 µL forward primer, 0.5 µL reverse primer,
and 1 µL DNA. The primers used for this study were
12S (12SZ-L: 5′-AAAGGTTTGGTCCTAGCCTT-3′ and
12SK-H: 5′ -TCCRGTAYRCTTACCDTGTTACGA-3′;
Goebel et al., 1999) and 16S, which included tRNAval
(12sm: 5′ -GGCAAGTCGTAACATGGTAAG-3′ and 16sd:
5′-CTCCGGTCTGAACTCAGATCACGTAG-3′; Pauly
et al., 2004; Oliver et al., 2015). Annealing temperature
for 12S and 16S was 52°C and 51°C, respectively. PCR
products were puried with ExoSAP-IT®. Sequencing was
performed by Macrogen (Netherland). Sequences were edited
and assembled in Geneious v 8.0 (Kearse et al., 2012) prior
to alignment.
We combined our 12S and 16S (including tRNAval) sequence
data (n = 9) with the Pulchrana dataset downloaded from
GenBank (n = 24), 22 of which came originally from Brown
& Siler (2013) and were later incorporated by Chan et al.
(2014), along with two from Matsui et al. (2012). These
downloaded sequences also comprise a species group of
related marsh frogs (i.e., P. banjarana, P. glandulosa, and P.
baramica) that we used as outgroup. Sequence information
and GenBank accession numbers are provided in Table 2.
We aligned sequences for each locus using MAFFT v7.7
(Katoh & Standley, 2013), as implemented in Geneious v 8.0
and concatenated the aligned sequences for a nal alignment
of 2,285 base pairs. Partition Finder V1.1.1 (Lanfear et al.,
2012) was used to determine the best partitioning schemes and
model of substitutions of the concatenated sequences under
the Bayesian Information Criterion (BIC) using the “greedy”
search algorithm. GTR+I+G was selected as the best model,
however, phylogenetic estimation using maximum likelihood
(ML) was performed with a GTR+G model because the
25 discrete categories are better at approximating invariant
sites (Stamatakis, 2014). The aforementioned analysis was
executed using RAxML v.8 (Stamatakis, 2006, 2014) at the
CIPRES Science Gateway server (Cyberinfrastructure for
Phylogenetic Research; www.phylo.org/sub.sections/portal;
Miller et al., 2010). We used MrBayes v3.2.6 (Huelsenbeck
& Ronquist, 2001; Ronquist & Huelsenbeck, 2003) with the
same server to perform Bayesian inference (BI) analysis using
default priors. The MCMC sampling was performed for 50
million generations using two independent runs, each with
four chains, and sampling every 1,000 generations with a 25%
burn-in. Successful convergence of runs was assessed using
trace plot and ESS values (>200) in Tracer v.1.6 (Rambaut &
Drummond, 2009). The output from RAxML and MrBayes
analyses was visualised in FigTree v1.4.3 (Rambaut, 2007).
Corresponding gures were prepared using CorelDRAW X6.
Bootstrap (BS) node values ≥ 70 (Hillis & Bull, 1993) and
a posterior probability (PP) of ≥ 0.95 (Alfaro et al., 2003;
Huelsenbeck & Rannala, 2004; Mulcahy et al., 2011) were
considered high support.
RESULTS
Phylogenetic analyses. Our final concatenated (12S +
16S + tRNAval) sequence matrix consisted of 2,285 bp and
included 5.32% gaps and undetermined character states. The
topology of the optimal ML tree (lnL = -12646.925261) was
identical to that inferred in our BI analysis (Fig. 1). These
topologies match the topology recovered by Brown & Siler
(2013) and Chan et al. (2014), except for the arrangement
of Pulchrana signata. Chan et al. (2014) suggested P.
signata as sister taxon to a four taxon clade comprising
([P. mangyanum + P. moellendorf] + [P. grandocula + P.
similis]). In our trees, P. signata was instead recovered as a
sister taxon to P. mangyanum + P. moellendorf. The clade
comprising these species was sister to P. grandocula + P.
similis. Nodal support was generally high for both BS and
PP, with exceptions including the node joining P. signata
to P. moellendorf + P. mangyanum (BS = 69; PP = 0.96),
the sister taxon relationship between P. grandocula and P.
similis (BS = 74; PP = 0.89), the node joining the two P.
similis (BS = 59; PP = 0.62), and the node joining the two P.
siberu from Pagai Selatan and Sipora (BS = 57; PP = 0.63).
The two individuals from Jambi formed a strongly supported
clade (BS = 100; PP = 1) together with the Pulchrana
centropeninsularis sample from Brown & Siler (2013),
with negligible genetic divergence (uncorrected p-distance
= 0.003; Table 3). There was strong support (BS = 98;
PP =1; Fig. 1) for the two individuals from Aceh being
nested within the samples from Sumatera Utara, previously
reported as being “Hylarana cf. siberu [Sumatra]” and “sp
Sumatra” by Brown & Siler and (2013) and Chan et al.
(2014) , respectively. All P. siberu samples formed a strongly
supported clade (BS = 100; PP = 1) with minimal genetic
divergence (Table 3) among the three sampled islands
(Siberut, Sipora and Pagai Selatan).
Comparison between individuals of Pulchrana
centropeninsularis from the Malay Peninsula and
Sumatra. The observed morphological character states in
three adult males from Hutan Harapan, Jambi Province,
MZB.Amph.28765 (S 02.18010°, E 103.50215°); MZB.
Amph.28766–67 (S 02.18431°, E 103.36633°) corroborate
our genetic results; the specimens can be assigned to P.
centropeninsularis following the characteristics described
in Chan et al. (2014). A comparison of morphometric
characters between P. centropeninsularis from the Malay
Peninsula and Jambi is shown in Table 4 (individual
measurements) and Table 5 (means and standard deviations).
We noticed minor differences between specimens from the
Malay Peninsula (Chan et al., 2014) and Jambi (Sumatra)
specimens, and provide the following enumeration of these
observations (with opposing character states for Sumatra
specimens in parentheses): dorsal skin texture smooth to
nely granulated (nely granulated to granular; Fig. 2); tear
drop-shaped choana (circular); NSD/END = 70.0% (NSD/
END = 44.4–55.7%); HG/BL = 52.5–57.5% (HG/BL =
31.6–45.9%); webbing formula for holotype (ZRC1.10536)
and paratype (DWNP 1189) (Chan et al., 2014): I(1/2 ― 2)
II(1 ― 21/2)III(1 ― 3)IV(3 ― 1)V [webbing formula for

281
RAFFLES BULLETIN OF ZOOLOGY 2018
Table 2. List of specimens included in our molecular phylogenetic analyses, corresponding museum catalog numbers, collection localities,
GenBank accession numbers, and original citations.
Species Voucher* Locality GenBank no. Citation
Pulchrana baramica FMNH 248218 Brunei, Borneo Island, Belait
District
KF477628 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana baramica KUHE 53640 Malaysia, Borneo Island,
Sarawak, Mulu
AB719217,
AB719234
Matsui et al., 2012
Pulchrana banjarana LSUHC 5128 Malaysia, Malay Peninsula,
Pahang, Cameron Highlands
KF477644 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana banjarana ZRC 8326 Malaysia, Malay Peninsula,
Pahang, Cameron Highlands
KF477645 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana centropeninsularis DWNP 0489 Malaysia, Malay Peninsula,
Pahang, Kuala Gandah
KF477745 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana centropeninsularis MZB.
Amph.28765
Indonesia, Sumatra Island,
Jambi Province, Harapan Rain
Forest
MG783353,
MG783362
This study
Pulchrana centropeninsularis MZB.
Amph.28766
Indonesia, Sumatra Island,
Jambi Province, Harapan Rain
Forest
MG783352,
MG783361
This study
Pulchrana fantastica, new
species
MZB.
Amph.28946
Indonesia, Sumatra Island, Aceh
Province, Gunung Leuseur
National Park
MG783359,
MG783367
This study
Pulchrana fantastica, new
species
MZB.
Amph.28891
Indonesia, Sumatra Island, Aceh
Province, Taman Buru Linge
Isaq
MG783360,
MG783368
This study
Pulchrana fantastica, new
species
MK 334 Indonesia, Sumatra Island,
Sumatera Utara Province,
Langkat, Bandar Baru, Batak
Mountains
KF477646 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana fantastica, new
species
MZB.
Amph.13011
(MK 335)
Indonesia, Sumatra Island,
Sumatera Utara Province,
Langkat, Bandar Baru, Batak
Mountains
KF477648 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana glandulosa KUHE 53618 Malaysia, Borneo Island,
Sarawak, Mulu
AB719206,
AB719223
Matsui et al., 2012
Pulchrana grandocula KU 306492 Philippines, Samar Island,
Samar Province, Municipality
of San Jose de Baun, Barangay
Poblacion
KF477660 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana grandocula PNM 8848 Philippines, Mindanao
Island, Davao City Province,
Municipality of Calinan,
Barangay Malagos
KF477676 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana mangyanum KU 303566 Philippines, Mindoro Island,
Municipality of Paypayama,
Barangay Carmundo
KF477687 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana mangyanum KU 303578 Philippines, Mindoro Island,
Municipality of Bongabong,
Barangay Formon
KF477686 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana moellendorf KU 309009 Philippines, Palawan Island,
Palawan Province, Municipality
of Puerto Princesa City,
Barangay Irawan
KF477696 Brown & Siler, 2013
and Chan et al., 2014

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Arin et al.: New Pulchrana from Sumatra
Species Voucher* Locality GenBank no. Citation
Pulchrana moellendorf KU 327050 Philippines, Palawan Island,
Palawan Province, Municipality
of Nara, Barangay Estrella Falls
KF477695 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana picturata FMNH 235707 Malaysia, Borneo Island, Sabah,
Kota Marudu
KF477729 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana picturata FMNH 238866 Malaysia, Borneo Island, Sabah,
Tenom District
KF477731 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana picturata FMNH 266930 Indonesia, Sumatra Island,
Sumatera Barat Province, Limau
Manis
KF477717 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana picturata FMNH 266944 Indonesia, Sumatra Island,
Sumatera Barat Province,
Payakumbuh
KF477701 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana siberu BJE 203 Indonesia, Siberut Island,
Sumatera Barat Province
KF477741 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana siberu BJE 236 Indonesia, Siberut Island,
Sumatera Barat Province
KF477743 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana siberu MVZ 272090 Indonesia, Siberut Island,
Sumatera Barat Province
MG783357,
MG783365
This study
Pulchrana siberu MVZ 272091 Indonesia, Siberut Island,
Sumatera Barat Province
MG783358,
MG783366
This study
Pulchrana siberu MVZ 272082 Indonesia, Pagai Selatan Island,
Sumatera Barat Province
MG783354, NA This study
Pulchrana siberu MVZ 272086 Indonesia, Pagai Selatan Island,
Sumatera Barat Province
MG783355,
MG783363
This study
Pulchrana siberu MVZ 272089 Indonesia, Sipora Island,
Sumatera Barat Province
MG783356,
MG783364
This study
Pulchrana signata FMNH 238842 Malaysia, Borneo Island, Sabah,
Sipitang District, Mendolong
KF477746 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana signata ZRC 1.12388 Malaysia, Borneo Island,
Sarawak, Matang
KF477748 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana similis TNHC 63007 Philippines, Luzon Island,
Camarines Norte Province,
Municipality of Naga City,
Barangay Panicuason
KF477764 Brown & Siler, 2013
and Chan et al., 2014
Pulchrana similis PNM 5536 Philippines, Luzon Island,
Laguna Province, Municipality
of Los Banos, University of
the Philippines campus, Mt.
Makiling
KF477776 Brown & Siler, 2013
and Chan et al., 2014
*FMNH = Field Museum of Natural History; LSUHC = La Sierra University Herpetological Collection; ZRC = Zoological Reference
Collection, Lee Kong Chian Natural History Museum, Singapore; MK = Mistar Kamsi eld number; DWNP = Department of Wildlife
and National Parks, Malaysia; BJE = Ben J. Evans eld number; KU = University of Kansas; PNM = Philippines National Museum;
TNHC = Texas National History Museum, University of Texas, Austin; MVZ = Museum of Vertebrate Zoology, University of California,
Berkeley; MZB = Museum Zoologicum Bogoriense, Indonesia.

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Fig. 1. Phylogenetic tree depicting the relationship between Pulchrana fantastica, new species and congeners based on BI. ML tree shows
similar topology as BI. Node support representing bootstraps value and posterior probabilities (BS/PP).
Fig. 2. Left to right; examples of dorsal skin textures in members of the Pulchrana siberu, P. centropeninsularis, and P. fantastica, new
species: (a) nely granulated (MZB.Amph.28896); (b) granular (MZB.Amph.28765); (c) or with white tipped keratinised asperities (MZB.
Amph.13011). Scale bar = 3 mm. Photo by G. Cahyadi.
MZB.Amph.28765: I(1- ― 1+)II(10 ― 2-)III(1- ― 20)IV(2- ―
1/2)V; for MZB.Amph.28766: I(1/2―1+)II(1/2―11/2)III(1-
― 20)IV(2- ― 1/2)V; for MZB.Amph.28767: I(1/2 ― 1+)
II(1/2 ― 11/2)III(1- ― 20)IV(11/2 ― 1/2)V]. We consider
these differences a representation of intraspecic variation.
In available literatures on Pulchrana centropeninsularis from
the Malay Peninsula (Leong & Lim, 2004; Chan & Norhayati,
2009; Chan et al., 2014), this species was reported as a
possible obligate swamp-adapted specialist. However, our
Jambi specimens were collected from a stream in a lowland
secondary forest (maximum elevation 50 m) that had been
selectively logged approximately 40 years ago (Fig. 3a).
The stream’s width was approximately 0.5–5.7 m, and depth
0.01–1 m with a slow ow. The water was silty and had low
visual clarity. In total, we encountered 14 specimens at the
site, of which 12 were found along the stream. The resting
sites of the specimens were recorded at 0.6–3.5 m distance
from the water, always in forest habitat, away from the stream
bank. The majority of these individuals perched on branches

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Table 3. Uncorrected p-distances calculated from 16S sequences (in MEGA v.7) between Pulchrana fantastica, new species and congeners.
1 2 3 4 5 6 7 8 9 10 11
1P. glandulosa KUHE 53618
2P. baramica KUHE 53640 0.064
3P. baramica FMNH 248218 0.064 0.005
4P. banjarana ZRC 8326 0.163 0.172 0.170
5P. banjarana LSUHC 5128 0.160 0.168 0.167 0.012
6P. picturata FMNH 235707 0.169 0.155 0.154 0.161 0.163
7P. picturata FMNH 238866 0.169 0.166 0.166 0.170 0.172 0.066
8P. picturata FMNH 266944 0.151 0.150 0.152 0.151 0.152 0.087 0.090
9P. picturata FMNH 266930 0.157 0.150 0.151 0.150 0.149 0.088 0.090 0.021
10 P. signata ZRC 112388 0.155 0.152 0.152 0.167 0.169 0.115 0.104 0.109 0.108
11 P. signata FMNH 238842 0.154 0.152 0.152 0.166 0.166 0.110 0.103 0.108 0.109 0.022
12 P. siberu MVZ 272086 0.156 0.156 0.153 0.147 0.145 0.125 0.131 0.128 0.124 0.141 0.133
13 P. siberu MVZ 272089 0.160 0.157 0.154 0.150 0.149 0.128 0.134 0.128 0.124 0.145 0.138
14 P. siberu MVZ 272090 0.155 0.154 0.151 0.150 0.148 0.129 0.135 0.129 0.121 0.141 0.138
15 P. siberu MVZ 272091 0.154 0.153 0.150 0.150 0.147 0.128 0.134 0.128 0.120 0.140 0.137
16 P. siberu BJE 202 0.154 0.153 0.150 0.151 0.147 0.129 0.135 0.130 0.122 0.141 0.138
17 P. siberu BJE 236 0.154 0.153 0.150 0.151 0.147 0.129 0.135 0.130 0.122 0.141 0.138
18 P. centropeninsularis MZB.
Amph.28765
0.160 0.154 0.153 0.145 0.146 0.123 0.136 0.122 0.123 0.131 0.126
19 P. centropeninsularis MZB.
Amph.28766
0.160 0.154 0.153 0.145 0.146 0.123 0.136 0.122 0.123 0.131 0.126
20 P. centropeninsularis DWNP
0489
0.160 0.154 0.153 0.143 0.144 0.123 0.136 0.122 0.123 0.130 0.124
21 P. fantastica, new species
MK334
0.155 0.155 0.152 0.150 0.145 0.127 0.130 0.140 0.136 0.130 0.131
22 P. fantastica, new species
MZB.Amph.13011
0.157 0.157 0.154 0.151 0.147 0.130 0.132 0.142 0.138 0.132 0.133
23 P. fantatisca, new species
MZB.Amph.28946
0.158 0.156 0.153 0.151 0.147 0.128 0.131 0.140 0.136 0.134 0.133
24 P. fantastica, new species
MZB.Amph.28891
0.158 0.156 0.153 0.150 0.145 0.128 0.131 0.138 0.134 0.134 0.133
25 P. moellendorf KU 327050 0.163 0.155 0.154 0.170 0.164 0.113 0.109 0.117 0.113 0.106 0.103
26 P. moellendorf KU 309009 0.163 0.155 0.154 0.170 0.164 0.113 0.109 0.117 0.113 0.106 0.103
27 P. mangyanum KU 303566 0.161 0.156 0.154 0.164 0.161 0.109 0.107 0.106 0.104 0.094 0.090
28 P. mangyanum KU 303578 0.162 0.157 0.155 0.165 0.162 0.109 0.107 0.106 0.104 0.093 0.091
29 P. grandocula KU 306492 0.173 0.172 0.174 0.171 0.171 0.112 0.114 0.114 0.118 0.110 0.109
30 P. grandocula PNM 8848 0.170 0.170 0.171 0.169 0.167 0.114 0.110 0.110 0.114 0.108 0.103
31 P. similis TNHC 63007 0.170 0.170 0.172 0.174 0.173 0.111 0.109 0.106 0.111 0.104 0.101
32 P. similis PNM 5536 0.171 0.171 0.173 0.172 0.171 0.110 0.110 0.109 0.113 0.107 0.102

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12 13 14 15 16 17 18 19 20 21
1P. glandulosa KUHE 53618
2P. baramica KUHE 53640
3P. baramica FMNH 248218
4P. banjarana ZRC 8326
5P. banjarana LSUHC 5128
6P. picturata FMNH 235707
7P. picturata FMNH 238866
8P. picturata FMNH 266944
9P. picturata FMNH 266930
10 P. signata ZRC 112388
11 P. signata FMNH 238842
12 P. siberu MVZ 272086
13 P. siberu MVZ 272089 0.010
14 P. siberu MVZ 272090 0.016 0.014
15 P. siberu MVZ 272091 0.015 0.015 0.003
16 P. siberu BJE 202 0.017 0.015 0.001 0.004
17 P. siberu BJE 236 0.017 0.015 0.001 0.004 0.000
18 P. centropeninsularis MZB.
Amph.28765 0.083 0.084 0.083 0.082 0.083 0.083
19 P. centropeninsularis MZB.
Amph.28766 0.083 0.084 0.083 0.082 0.083 0.083 0.000
20 P. centropeninsularis DWNP 0489 0.082 0.083 0.083 0.082 0.083 0.083 0.003 0.003
21 P. fantastica, new species MK334 0.100 0.103 0.101 0.100 0.101 0.101 0.103 0.103 0.103
22 P. fantastica, new species MZB.
Amph.13011 0.102 0.105 0.103 0.102 0.103 0.103 0.103 0.103 0.103 0.004
23 P. fantastica, new species MZB.
Amph.28946 0.102 0.105 0.105 0.104 0.105 0.105 0.102 0.102 0.102 0.011
24 P. fantastica, new species MZB.
Amph.28891 0.100 0.103 0.103 0.102 0.103 0.103 0.102 0.102 0.102 0.013
25 P. moellendorf KU 327050 0.131 0.132 0.135 0.134 0.135 0.135 0.145 0.145 0.143 0.137
26 P. moellendorf KU 309009 0.131 0.132 0.135 0.134 0.135 0.135 0.145 0.145 0.143 0.137
27 P. mangyanum KU 303566 0.125 0.128 0.131 0.130 0.131 0.131 0.130 0.130 0.129 0.133
28 P. mangyanum KU 303578 0.125 0.128 0.131 0.130 0.131 0.131 0.131 0.131 0.130 0.133
29 P. grandocula KU 306492 0.132 0.134 0.133 0.132 0.133 0.133 0.140 0.140 0.138 0.141
30 P. grandocula PNM 8848 0.130 0.132 0.134 0.133 0.134 0.134 0.135 0.135 0.133 0.142
31 P. similis TNHC 63007 0.132 0.134 0.136 0.135 0.136 0.136 0.138 0.138 0.136 0.137
32 P. similis PNM 5536 0.130 0.132 0.134 0.133 0.134 0.134 0.136 0.136 0.134 0.140

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22 23 24 25 26 27 28 29 30 31
1P. glandulosa KUHE 53618
2P. baramica KUHE 53640
3P. baramica FMNH 248218
4P. banjarana ZRC 8326
5P. banjarana LSUHC 5128
6P. picturata FMNH 235707
7P. picturata FMNH 238866
8P. picturata FMNH 266944
9P. picturata FMNH 266930
10 P. signata ZRC 112388
11 P. signata FMNH 238842
12 P. siberu MVZ 272086
13 P. siberu MVZ 272089
14 P. siberu MVZ 272090
15 P. siberu MVZ 272091
16 P. siberu BJE 202
17 P. siberu BJE 236
18 P. centropeninsularis MZB.Amph.28765
19 P. centropeninsularis MZB.Amph.28766
20 P. centropeninsularis DWNP0489
21 P. fantastica, new species MK334
22 P. fantastica, new species MZB.
Amph.13011
23 P. fantastica, new species MZB.
Amph.28946 0.013
24 P. fantastica, new species MZB.
Amph.28891 0.015 0.002
25 P. moellendorf KU 327050 0.140 0.134 0.132
26 P. moellendorf KU 309009 0.140 0.134 0.132 0.000
27 P. mangyanum KU 303566 0.133 0.133 0.133 0.058 0.058
28 P. mangyanum KU 303578 0.132 0.133 0.133 0.058 0.058 0.002
29 P. grandocula KU 306492 0.142 0.143 0.142 0.103 0.103 0.096 0.096
30 P. grandocula PNM 8848 0.143 0.144 0.143 0.098 0.098 0.090 0.090 0.030
31 P. similis TNHC 63007 0.138 0.139 0.140 0.099 0.099 0.098 0.098 0.024 0.025
32 P. similis PNM 5536 0.141 0.142 0.143 0.099 0.099 0.091 0.091 0.024 0.017 0.011

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Fig. 3. Habitat of Pulchrana centropeninsularis, Hutan Harapan, Jambi (a) and P. fantastica, new species, Taman Buru Linge Isaq, Aceh
(b, c). Photo by A. Jankowski (a); G. Cahyadi (b, c).

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Table 4. Morphometric variation (in mm) between specimens of Pulchrana centropeninsularis from the Malay Peninsula and Jambi.
P. centropeninsularis
(Malay Peninsula; Chan et al., 2014)
P. centropeninsularis
(Jambi; this study)
ZRC1.10536
(holotype)
DWNP1189
(paratype) MZB.Amph.28765 MZB.Amph.28766 MZB.Amph.28767
SVL 37.4 37.6 40.2 35.5 40.4
HL 14 15.2 15.2 14.5 15.2
HW 12 12.5 10.3 10.4 11.3
SL 5.9 6.4 6.2 5.8 6.2
IOD 3.2 3.5 3.7 3.9 3.8
IND 3.7 3.6 3.1 3.3 3.5
ED 6.0 5.1 5.3 5.4 5.3
TD 4.8 2.2 3.1 3.2 3.4
BL 7.3 8.0 8.9 9.1 8.8
FAL 8.1 8.2 9.6 9.2 10.5
FL 17.6 17.7 17.9 17.5 19.9
TBL 19.9 19.2 20.4 20.3 21.3
TL 12.9 10.6 11.0 11.1 11.6
HG 4.2 4.2 3.3 2.9 4.1
Table 5. Summary statistics of morphometric data for Pulchrana centropeninsularis, P. siberu, and P. fantastica, new species. Table
entries are Mean (± SD), followed by range, in mm.
P. centropeninsularis P. centropeninsularis P. siberu P. fantastica, new species
Malay Peninsula, n=2;
Chan et al., 2014 Jambi, n = 3; this study
Siberut Island, n = 3;
Brown & Guttman 2002;
Chan et al., 2014
Aceh and Sumatera Utara,
n = 12; this study
SVL 37.5 ± 0.1 (37.4–37.6) 38.7 ± 2.7 (35.5–40.4) 37.0 ± 2.2 (35.4–39.5) 42.4 ± 1.3 (40.3–45.2)
HL 14.6 ± 0.8 (14.0–15.2) 14.8 ± 0.4 (14.5–15.2) 15.7 ± 0.1 (15.6–15.9) 15.0 ± 0.7 (14.1–16.0)
HW 12.3 ± 0.4 (12.0–12.5) 10.6 ± 0.6 (10.3–11.3) 13.0 ± 0.3 (12.6–13.2) 11.5 ± 0.9 (9.5–12.4)
SL 6.2 ± 0.4 (5.9–6.4) 6.0 ± 0.2 (5.8–6.2) 7.0 ± 0.5 (6.5–7.4) 6.7 ± 0.2 (6.4–7.0)
IOD 3.4 ± 0.2 (3.2–3.5) 3.8 ± 0.1 (3.7–3.9) 4.2 ± 0.3 (3.8–4.5) 4.0 ± 0.3 (3.4–4.4)
IND 3.7 ± 0.1 (3.6–3.7) 3.3 ± 0.2 (3.1–3.5) 4.1 ± 0.2 (4.0–4.3) 4.0 ± 0.2 (3.5–4.3)
ED 5.6 ± 0.6 (5.1–6.0) 5.4 ± 0.1 (5.3–5.4) 5.3 ± 0.3 (5.1–5.3) 6.0 ± 0.3 (5.5–6.5)
TD 3.5 ± 1.8 (2.2–4.8) 3.2 ± 0.2 (3.1–3.4) 3.6 ± 0.3 (3.3–3.9) 3.5 ± 0.2 (3.2–3.8)
BL 7.7 ± 0.5 (7.3–8.0) 8.9 ± 0.1 (8.8–9.1) 8.0 ± 0.6 (7.3–8.4) 10.1 ± 0.5 (9.1–10.6)
FAL 8.2 ± 0.1 (8.1–8.2) 9.8 ± 0.6 (9.2–10.5) 9.7 ± 0.5 (9.2–10.1) 9.5 ± 0.6 (8.5–10.5)
FL 17.7 ± 0.1 (17.6–17.7) 18.4 ± 1.3 (17.5–19.9) 18.7 ± 1.6 (17.0–20.2) 21.5 ± 0.9 (20.0–23.5)
TBL 19.6 ± 0.5 (19.2–19.9) 20.6 ± 0.5 (20.3–21.3) 20.9 ± 0.8 (20.0–21.6) 23.2 ± 0.9 (22.1–25.0)
TL 11.8 ± 1.6 (10.6–12.9) 11.2 ± 0.3 (11.0–11.6) 12.1 ± 0.4 (11.7–12.5) 12.3 ± 0.8 (11.2–13.9)
HG 4.2 ± 0.0 (4.2–4.2) 3.4 ± 0.6 (2.9–4.1) 4.5 ± 0.3 (4.3–4.8) 3.9 ± 0.3 (3.2–4.3)
or leaves of small bushes or saplings at 0.2–0.5 m above
ground. Male advertisements calls were heard at these resting
sites, suggesting the stream may have been the reproduction
site of P. centropeninsularis. Pulchrana picturata was
recorded in sympatry with P. centropeninsularis at that site.
We also recorded P. centropeninsularis at other localities
in Jambi with similar habitat types (Fig. 4).
TAXONOMY
Based on the phylogenetic placement and morphological
distinctness of the Sumatran lineage, we consider these
specimens to represent a new species, exhibiting characteristics
of a separately evolving lineage, in accordance with the
General Unied Lineage-based concept (sensu de Queiroz,
2005). As a result, we describe this unnamed North Sumatran
species as a member of the Pulchrana signata Complex,
within which it is unequivocally nested (Brown & Siler,
2013; Chan et al., 2014), and of which it exhibits diagnostic
characters as formalised by Brown & Guttman (2002).

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Fig. 4. Geographical distribution of Pulchrana centropeninsularis (white circles), P. fantastica, new species (black triangles), and P. siberu
(red stars). Locality information: Mane (1); Taman Buru Linge Isaq (2); Taman Nasional Gunung Leuser (3); Bandar Baru (4), Lakum
Forest Reserve & Kuala Gandah, Pahang (5); Bukit Tigapuluh (6); Hutan Harapan (7); Siberut (8); Sipora (9); and Pagai Selatan (10).
Type locality for each species indicated by arrow. Map was prepared using GeoMapApp (Ryan et al., 2009).
Ranidae Batsch, 1796
Pulchrana Dubois, 1992
Pulchrana fantastica, new species
Splendid Stream Frogs (recommended common English
name); Katak Elok (Bahasa Indonesia)
(Fig. 5a, b)
Hylarana cf. siberu (Montane NW Sumatra [Brown & Siler, 2013])
Hylarana sp. Sumatra (Chan et al., 2014)
Holotype. MZB.Amph.28891 (adult male, Fig. 6a–d) from
Taman Buru Linge Isaq, Aceh Province, Sumatra, Indonesia
(N 04.35868° E 097.24404°, 450 m elevation), coll. U. Arin
and G. Cahyadi, 7 March 2014 at 2054 hours.
Paratypes (6). MZB.Amph.28898 (adult male) and MZB.
Amph.28890 (adult male), same information as the holotype,
collected between 2036–2145 hours; MZB.Amph.28892–93,
MZB.Amph.28894 (Fig. 5a) and MZB.Amph.28896 (adult
males), at N 04.37958° E 097.29158° 1,000 m elevation, 9
March 2014, coll. U. Arin and G. Cahyadi.
Referenced specimens (11). Seven specimens were
collected from Taman Buru Linge Isaq, Aceh Province
by U. Arin and G. Cahyadi: MZB.Amph.28889 (male),
MZB.Amph.28943 (juvenile; Fig. 5b), MZB.Amph.28948
(juvenile), at N 04.35868° E 097.24404°, 450 m elevation,
7 March 2014; MZB.Amph.28945 (juvenile) and MZB.
Amph.28947 (juvenile), at N 04.36018° E 097.24580°, 450
m elevation, 7 March 2014; MZB.Amph.28897 (male), at
N 04.338036° E 097.28096°, 600 m elevation, 8 March
2014; and MZB.Amph.28944 (juvenile), at N 04.37958°
E 097.29158°, 1,000 m elevation, 9 March 2014. MZB.

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Fig. 5. Plates comparing the three closely allied Sumatran Pulchrana species (a) P. fantastica, new species, MZB.Amph.28894, adult
male, paratype, Aceh; (b) P. fantastica, new species, MZB.Amph.28943, juvenile, Aceh; (c) P. centropeninsularis, adult male, Jambi; (d)
P. siberu, female, Pagai Selatan. Photo by U. Arin (a, b, d); A. Jankowski (c).
Amph.28895 (male), at a locally protected forest in Mane,
Kecamatan Ulu Masen (N 04.89949° E 096.13168°, 700 m
elevation), 21 March 2014, coll. U. Arin and G. Cahyadi.
MZB.Amph.28946 (juvenile), at Marpunge, Taman Nasional
Gunung Leuser, Aceh Province (N 03.77146° E 097.63773°,
1,065 m elevation), 23 February 2014, coll. U. Arin and
G. Cahyadi. MZB.Amph.13015 and MZB.Amph.13011,
both males, at Bandar Baru, Sumatera Utara Province (N
03.26287°, E 098.46793°), 5 December 2006 and 10 January
2007, coll. M. Kamsi.
Diagnosis. The following unique combinations of characters
distinguish Pulchrana fantastica, new species, from its
congeners: (1) a medium size frog, SVL adult males (n =
12) 40.3–45.2 mm; (2) males with large humeral glands
(3.2–4.3 mm) on anteroventral surface of brachium, paired
internal subgular vocal sacs, nuptial pads absent; (3) dorsal
skin nely granular to granulated, with or without keratinised
white asperities at tip of each granule (Fig. 2); (4) webbing
formula: I(1 ― 1–11/2)II(1/2 ― 2)III(1 ― 2–21/3)IV(2–
21/3 ― 1)V; (5) dorsolateral stripe, thin (0.7–0.9 mm),
orange, continuous, anteriorly confluent and posteriorly
interconnected by spots; (6) middorsum black with orange
line or combination of spots and line in the center, variable
in number and length of the line and spots, black without
marking in juveniles; (7) dense spots on anks and dorsal
surface limbs, cream or yellow to orange, shape of spots
elongated or circular, variable in size; (8) skin of venter
smooth, greyish or brown with small light dots on throat and
chest, occasionally extending posteriorly to abdomen; (9) iris
background black, dense orange stippling ventrally, orange
reticulation dorsally, golden centrally, with orange-golden
line encircling pupil; (10) upper and lower lip grey or brown
with cream or yellow spots (upper lip: 3–7; lower lip: 2–5).
Description of holotype. Adult male with large humeral
gland (HG/BL = 40.6%) on anteroventral surface of brachium,
paired internal subgular vocal sacs, nuptial pad absent; body
slender; head longer than wide (HL/HW = 129.8%); snout
obtusely pointed in dorsal view, slightly protruding in lateral
view; nares closer to snout than to eye (NSD/END = 56.8%);
canthus rostralis sharp, constricted behind nares; loreal region
sloping, deeply concave; vomerine teeth distinct, between
choana, left (n=3) and right (n=2), teeth barely separated
(distance 1.0 mm); choana circular (diameter = 1.0 mm),
interchoanal distance 5.3 mm; tongue lanceolate, widening
posteriorly, deeply notched in the center, 17.3% free of
its total length; eye diameter > interorbital distance (ED/
IOD = 151.2%); internarial distance subequal interorbital
distance (IND/IOD = 95.3%); tympanum diameter 58.5%
eye diameter; supratympanic fold conspicuous.

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Dorsum granulated (Fig. 2); anks nely granular; venter
smooth; forelimb relatively slender. Forearm length subequal
to brachial length (BL/FAL = 103.9%); ngers long and
slender, without webbing; Fin1L/Fin2L = 103.8%, Fin1L/
Fin4L = 86.5%, Finger III longest; fingertips slightly
expanded into rounded disc, circummarginal groove present;
disc width of nger wider than width of terminal phalanx
of nger: Fin1DW/Fin1TPW = 142.9%, Fin2DW/Fin2TPW
= Fin3DW/Fin3TPW = Fin4DW/Fin4TPW = 128.6%;
subarticular tubercles present, round, raised prominently;
one subarticular tubercle on Finger I and II, two on Finger
III and IV; supernumerary tubercles between the base of
each nger and subarticular tubercle present, smaller and
less prominent than subarticular tubercles, translucent; outer
metacarpal tubercle and palmar tubercle distinct, elongate
(OMCL/OMCW = 254.5%, PTL/PTW = 227.3%), in contact,
not prominent, subequal in length and width (OMCL/PTL =
112.0%, OMCW/PTW = 100.0%); inner metacarpal tubercle
oval, prominent, slightly longer than outer metacarpal tubercle
and palmar tubercle (IMCL/OMCL = 107.1%; IMCL/PTL
= 120.0%).
Hindlimbs long, tibia longer than femur (TBL/FL = 109.3%);
relative length of femur, tibia, and tarsus, to SVL is 50.2%,
54.9%, and 30.4%, respectively; skin texture of dorsal side
of tibia and posterior region of the thigh nely granulated;
tip of toes expanded, circummarginal groove present, widths
of the toe disc larger than widths of terminal phalanx of the
respective toes: Toe1DW/Toe1TPW = 118.2%, Toe2DW/
Toe2TPW = 137.5%, Toe3DW/Toe3TPW = Toe4DW/
Toe4TPW = 128.6%, Toe5DW/Toe5TPW = 133.3%;
subarticular tubercles distinct, round, highly elevated,
translucent; number of subarticular tubercle for each toe: I(1),
II(1), III(2), IV(3), V(2); relative toe length: I<II<V≤III<IV,
Toe3L/Toe5L = 104.4%; outer metatarsal tubercle raised,
oval (OMTL/OMTW = 112.5%), translucent; inner metatarsal
tubercle distinct and long (IMTL/IMTW = 200.0%), elevated,
translucent, larger than outer metatarsal tubercles (IMTL/
OMTL = 200.0%); webbing formula: I(1- ― 10)II(1/2 ― 2-)
III(1- ― 2+)IV(2- ― 1-)V.
Colouration. In life, dorsal skin black; middorsum with
orange spot behind the eyes continued by orange line
(divided into two, equal length) up to approximately level
of sacral vertebra, two yellow spots, and one orange spot in
the pelvic region (in the middle of two ilium, in line with
joint between ilium and femur); continuous straight, orange
stripes (width = 0.7 mm), from tip of snout, along canthus
rostralis, following lateral margin of palpebra, continuing
dorsolaterally to sacrum, breaking up into five orange-
round spots at the posterior pelvic region that connect the
dorsolateral stripes from both sides into a loop; anks brown,
lighter ventrally; yellow spots from behind tympanum to
groin; round cream spot present between tympanum and
eye; two cream spots at end of rictus; venter greyish-brown;
whitish dots on throat, chest, and abdomen; iris background
black, dense orange reticulation ventrally, orange reticulation
dorsally, with golden centrally, solid orange line encircling
pupil; upper lip brownish-grey with white spots (n = 6 on
left, 5 on right); lower labial region grey, with three small
white spots on each side of lower lip; dorsal surface of limbs
brown, with dense round spots and elongate, yellow to orange
markings, variable in size; small whitish spots on rear of
thigh; interdigital webbing brown. In preservative, dorsal
skin dark brown; anks and dorsal surface of limbs brown,
lighter than dorsum; ventral skin creamy brown, with whitish
spots; dorsolateral stripe and spots on dorsum, anks, and
dorsal surface of limbs faded to cream or whitish; iris grey.
Variation. We observed variation within 18 specimens
of Pulchrana fantastica, new species; comparison of
morphological traits among the seven type specimens of
P. fantastica, new species, is provided in Table 6. Dorsum
texture of adults nely granulated (at surface with distinct
granules; Fig. 2a) or granular (granule distinct, more
raised, with white tipped asperities present or absent; Fig.
2b, c). Juvenile specimens lack middorsal marking, adults
middorsum with markings (yellow line and or spots from
central postocular region, extending posteriorly to vent,
variable in length), except for MZB.Amph.28896 without
marking; in life, juvenile colouration of spots on dorsal
surfaces of limbs whitish or pale yellow (except for MZB.
Amph.28943 and MZB.Amph.28946 possess few orange
spots); venter grey to brown, with light dots, variably from
throat to abdomen; orange dorsolateral stripe in juveniles
and adults, straight, in most cases continuous from rostrum
to beyond sacrum, occasionally with one or two interruptions
of the stripe (Table 6); pattern of spots on dorsal surfaces
of limbs in adults vary in colour and shape: usually yellow
to orange, round or elongated, from two or more connected
spots; posterior surfaces of thighs brown (similar to dorsum),
with yellowish spots, smaller than those on the dorsum;
number of vomerine teeth 2–3 on each side; number of
spots on upper lip (left, right): 3–6, 3–7; number of spots on
lower lip 2–5 on each side; webbing formula I(1 ― 1–11/2)
II(1/2–1 ― 2)III(1 ― 2–21/3)IV(2–21/3 ― 1)V.
Range. Pulchrana fantastica, new species, is currently known
from Aceh Province (Marpunge, Taman Nasional Gunung
Leuser; Taman Buru Linge Isaq; Mane) and Sumatera Utara
Province (Bandar Baru, Langkat) at an elevation between
450–1,065 m (Fig. 4).
Natural history. The new species is currently known only
from primary forest. All Aceh specimens were collected from
within protected areas (Taman Nasional Gunung Leuser,
Taman Buru Linge Isaq, and local protected forest in Mane).
The holotype was rst observed calling from among leaf
litter, under low vegetation, about 2.5 m from a small stream
(2–3 m width). Two of the paratypes (MZB.Amph.28890
and MZB.Amph.28898) and three juveniles (referenced
specimens: MZB.Amph.28889, MZB.Amph.28943, MZB.
Amph.28948) were collected the same night at the type
locality. MZB.Amph.28889 was perched on a fern growing
over above dead log, approximately 1.0 m from a nearby
stream at 2038 hours. MZB.Amph.28890 was perched on
a dead branch in a stream, approximately 5 cm above the
surface of the water at 2145 hours. MZB.Amph.28898
was encountered at 2048 hours on a dead log (d = 30 cm),
approximately 1.2 m from the stream. MZB.Amph.28943

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Arin et al.: New Pulchrana from Sumatra
Table 6. Comparison of morphological traits between the type specimens of Pulchrana fantastica, new species.
Trait MZB.Amph.28891
(holotype)
MZB.Amph.28890
(paratype)
MZB.Amph.28892
(paratype)
MZB.Amph.28893
(paratype)
MZB.Amph.28894
(paratype)
MZB.Amph.28896
(paratype)
MZB.Amph.28898
(paratype)
SVL (mm) 42.8 41.8 41.1 40.3 42.9 43.2 42.3
HL (mm) 15.7 14.4 14.9 14.1 15.3 15.7 15.9
HW (mm) 12.1 11.3 11.6 11.1 12.1 12.3 12.2
SL (mm) 6.9 6.8 6.7 7.0 6.5 6.8 6.8
IOD (mm) 4.3 4.0 4.0 4.0 4.0 4.2 3.9
IND (mm) 4.1 4.0 3.9 4.3 4.3 4.1 4.1
ED (mm) 6.5 5.8 5.6 6.3 5.8 6.1 5.9
TD (mm) 3.8 3.2 3.6 3.5 3.5 3.4 3.2
BL (mm) 10.6 10.6 9.4 10.2 9.8 10.1 10.1
FAL (mm) 10.2 9.2 8.5 8.9 9.0 9.1 9.9
FL (mm) 21.5 21.2 20.9 21.6 22.2 21.7 21.7
TBL (mm) 23.5 22.9 22.2 22.8 22.3 22.9 23.7
TL (mm) 13.0 12.1 11.2 11.7 11.3 12.2 13.3
HG (mm) 4.3 4.1 3.6 3.2 3.5 4.1 3.7
TED (mm) 1.6 1.8 1.7 1.0 1.2 1.6 1.4
NSD (mm) 2.5 2.5 2.6 2.7 2.2 2.6 2.5
END (mm) 4.4 4.3 4.1 4.3 4.3 4.2 4.4
EED (mm) 7.9 7.5 7.2 7.5 7.2 7.5 8.0
DLSW (mm) 0.7 0.8 0.8 0.7 0.7 0.8 0.7
Toe1L (mm) 8.0 7.2 7.6 7.5 7.4 7.3 7.8
Toe2L (mm) 11.5 10.8 10.8 10.4 10.6 10.3 11.5
Toe3L (mm) 16.5 14.9 15.0 15.0 15.9 14.5 15.9
Toe4L (mm) 22.0 20.8 20.3 20.0 20.3 20.2 20.2
Toe5L (mm) 15.8 15.0 14.6 14.3 14.7 14.6 15.2
Toe1DW (mm) 1.3 1.1 1.0 1.2 1.1 1.1 0.9
Toe2DW (mm) 1.1 1.0 0.9 1.1 1.2 1.0 1.2
Toe3DW (mm) 0.9 1.1 1.1 1.1 1.3 1.0 1.2
Toe4DW (mm) 0.9 0.9 1.0 1.0 1.1 1.0 1.1
Toe5DW (mm) 0.8 0.8 0.9 0.9 1.2 0.7 1.0
Toe1TPW (mm) 1.1 0.8 0.6 0.7 0.6 0.7 0.5

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Trait MZB.Amph.28891
(holotype)
MZB.Amph.28890
(paratype)
MZB.Amph.28892
(paratype)
MZB.Amph.28893
(paratype)
MZB.Amph.28894
(paratype)
MZB.Amph.28896
(paratype)
MZB.Amph.28898
(paratype)
Toe2TPW (mm) 0.8 0.7 0.7 0.8 0.7 0.6 0.7
Toe3TPW (mm) 0.7 0.8 0.5 0.6 0.6 0.7 0.6
Toe4TPW (mm) 0.7 0.6 0.6 0.8 0.8 0.6 0.6
Toe5TPW (mm) 0.6 0.6 0.6 0.6 0.6 0.5 0.7
Fin1L (mm) 10.9 9.4 9.2 9.3 9.5 9.9 10.0
Fin2L (mm) 10.5 9.3 9.0 9.2 9.0 9.1 9.6
Fin3L (mm) 13.1 12.4 11.3 11.5 12.0 11.9 12.5
Fin4L (mm) 12.6 10.8 9.9 10.4 10.1 10.1 11.6
Fin1DW (mm) 1.0 1.0 1.0 1.1 1.2 1.0 1.0
Fin2DW (mm) 0.9 1.0 0.8 1.0 1.1 0.8 1.0
Fin3DW (mm) 0.9 1.2 0.8 1.0 1.2 1.0 1.1
Fin4DW (mm) 0.9 1.3 1.2 1.1 1.2 1.2 1.2
Fin1TPW (mm) 0.7 0.7 0.7 0.7 0.7 0.7 0.6
Fin2TPW (mm) 0.7 0.7 0.6 0.5 0.6 0.7 0.7
Fin3TPW (mm) 0.7 0.7 0.6 0.6 0.7 0.7 0.6
Fin4TPW (mm) 0.7 0.7 0.6 0.6 0.7 0.6 0.7
IMTL (mm) 1.8 1.7 1.8 1.7 2.0 1.9 1.8
OMTL (mm) 0.9 0.9 0.8 0.9 1.0 0.9 0.9
IMTW (mm) 0.9 0.8 0.9 0.9 1.0 1.1 1.0
OMTW (mm) 0.8 1.0 0.8 0.8 0.9 1.0 0.8
IMCL (mm) 3.0 2.3 2.1 2.3 2.3 2.1 2.7
OMCL (mm) 2.8 2.4 1.6 2.0 2.3 2.0 2.6
IMCW (mm) 1.6 1.2 1.3 1.6 1.3 1.7 1.5
OMCW (mm) 1.1 1.1 0.9 1.1 1.2 1.1 1.1
PTL (mm) 2.5 1.8 1.5 2.0 1.7 2.1 2.1
PTW (mm) 1.1 1.1 0.9 1.0 1.2 1.2 1.2

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Arin et al.: New Pulchrana from Sumatra
Trait MZB.Amph.28891
(holotype)
MZB.Amph.28890
(paratype)
MZB.Amph.28892
(paratype)
MZB.Amph.28893
(paratype)
MZB.Amph.28894
(paratype)
MZB.Amph.28896
(paratype)
MZB.Amph.28898
(paratype)
Number of spots on
upper labial (left,
right)
6, 5 4, 4 5, 5 4, 5 4, 4 4, 4 4, 7
Number of spots on
lower labial (left,
right)
3, 3 2, 2 3, 4 5, 3 4, 4 4, 4 5, 3
Number of vomerine
teeth (left, right) 3, 2 3, 3 3, 2 3, 3 3, 2 3, 3 3, 3
Dorsum texture granular without
white asperities
granular with white
asperities
granular without
white asperities
granular without
white asperities
granular without
white asperities nely granulated granular with white
asperities
Middorsum color
pattern (in life)
black with yellow
lines and spots
black with yellow
line and spots
black with yellow
line
black with yellow
spots
black with yellow
line black, unmarked black with yellow
line and spots
Dorsolateral stripes
shape
continuous, with one
left side interruption;
ve posterior
interconnecting spots
continuous;
four posterior
interconnecting spots
continuous;
three posterior
interconnecting spots
continuous, with
interruptions (right:
2, left: 1); three
interconnecting spots
continuous;
three posterior
interconnecting spots
continuous, with one
left side interruption;
each followed by
one spot posteriorly
continuous; one
interconnecting spot
Throat coloration grey with light spots brown with light
spots
brown with light
spots
brown with light
spots
brown with light
spots
greyish brown with
light spots grey with light spots
Venter coloration grey with light spots
reaching abdomen
brown with light
spots reaching
abdomen
brownish with
light spots reaching
abdomen
greyish brown with
light spots reaching
abdomen
brown with light
spots reaching
abdomen
greyish brown with
light spots reaching
abdomen
grey with light spots
reaching abdomen
Webbing formula
I(1- ― 10)II(1/2
― 2-)III(1- ― 2+)
IV(2- ― 1-)V
I(10 ― 11/2)II(1/2
― 2-)III(1- ― 2+)
IV(2+ ― 1-)V
I(1- ― 10)II(1/2
― 2-)III(1- ― 2+)
IV(2- ― 1-)V
I(10 ― 11/2)II(1/2
― 2-)III(1- ― 2+)
IV(2+ ― 1-)V
I(1- ― 10)II(1/2 ―
2-)III(1- ― 2+)IV(2+
― 1-)V
I(10 ― 11/2)II(1/2
― 2-)III(1- ― 21/3)
IV(2+ ― 1-)V
I(1- ― 10)II(1/2 ―
2-)III(1- ― 21/3)
IV(2- ― 1-)V

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was caught seated on an orchid leaf, on the forest oor
at 2036 hours. MZB.Amph.28948 was collected at 2036
hours from an orchid leaf, approximately 10 cm above the
ground. At the time, the nearby stream was narrower than
its usual width, due to low seasonal precipitation. Other
species recorded in the same vicinity included one species
of ranid frog (Huia sumatrana), two species of colubrid
snakes (Boiga nigriceps and B. jaspidea), one species of
scincid lizard (Eutropis sp.), and orangutans (Pongo abelii).
The forest type was a typical lowland dipterocap forest.
The four remaining paratypes (MZB.Amph.28892, MZB.
Amph.28894, MZB.Amph.28896, and MZB.Amph.28944)
were collected in the same region, but at higher elevation
(1,000 m). The stream at this elevation was surrounded by
primary forest, and was approximately 5–6 m wide. Large
rocks were prevalent and the stream water was silty and red
in colour. The resting perches of collected animals included
rocks, dead logs, and roots. Pictures of habitat for this species
are provided (Fig. 3b, c).
Etymology. The specic epithet is nominative feminine
derivative of the Greek phantastikós. We apply this
adjective with a contemporary spelling and an implied
meaning of “being beyond imagination” with reference to
the extraordinary beauty of this species.
Comparisons. Pulchrana banjarana, P. centropeninsularis,
P. fantastica, new species, P. guttmani, P. grandocula, P.
mangyanum, P. moellendorf, P. picturata, P. siberu, P.
signata, and P. similis can be distinguished from P. baramica,
P. glandulosa, P. laterimaculata, P. melanomenta, P. rawa,
and P. debussyi by having light spots (yellowish white,
grey, orange or red in life) on dark (often black) dorsum;
dorsolateral stripes present or absent, when present then
in the form of a continuous or broken stripe from snout to
sacral region or beyond.
Pulchrana centropeninsularis, P. fantastica, new species,
and P. siberu differ from P. banjarana, P. guttmani, P.
grandocula, P. mangyanum, P. moellendorf, P. picturata,
P. signata, and P. similis by the absence (vs. presence) of
nuptial pads in males; the presence of distinct pale spots on
the limbs (vs. broad bars or indistinct blotches in all other
species); the presence of orange to red dorsolateral stripes
(vs. white, yellow, pale orange or tan in other species), by
reduction in webbing of toes: one phalanx free for Toe III
and Toe V, and two to two and half phalanges free for Toe
IV (vs. web nearly complete) (Brown & Guttman, 2002).
Pulchrana fantastica, new species, (n = 12) can be
distinguished from P. centropeninsularis (Jambi specimens,
n males = 3) by larger body size (mean SVL 42.4 mm vs.
38.7 mm); larger humeral gland (mean HG length 3.9 mm
vs. 3.4 mm); number of spots on upper lip 3–7 (vs. 2–4)
on each side; number of spots on lower lip 2–5 (vs. 1–3)
on each side; mean ratio of tongue length of notched region
and total tongue length 18.5% (vs. 22.2%); dorsal skin
texture nely granulated to granular (vs. granular); with or
without (vs. without) white tipped asperity at center of each
granule; dorsolateral stripe thin, mean of width 0.8 mm (vs.
1.2 mm); middorsum black, marked with short or longer line
with breaks in adults and unmarked dorsum in juveniles (vs.
black, unmarked); ventral skin grey to brown, with white
spots on throat, chest, and sometimes to abdomen (vs. grey
to brown, with light spots on throat and light reticulation on
venter); mean of tibia length 23.2 mm (vs. 20.6 mm); yellow
to orange (vs. yellow spots), round or elongate (vs. usually
round), dense spots on dorsal side of limbs; webbing formula:
I(1 ― 1–11/2)II(1/2 ― 2)III(1 ― 2–21/3)IV(2–21/3 ― 1)
V [vs. I(1/2–1 ― 1–2)II(1/2–1 ― 11/2–21/2)III(1 ― 2–3)
IV(11/2–3 ― 1/2–1)V]. Morphological comparison showing
dorsal, ventral, palmar and plantar views of these two species
are provided in Fig. 6.
Pulchrana fantastica, new species, differs in morphology
from P. siberu (Dring et al., 1990; Brown & Guttman, 2002)
by larger body size (mean SVL 42.4 mm vs. 37.0 mm);
shorter humeral gland (mean HG length 3.9 mm vs. 4.5
mm); dorsal skin texture nely granulated to granular (vs.
granular); with or without white tipped asperity at center of
each granule; dorsolateral stripe thin, mean of width 0.8 mm
(vs. 1.1 mm in P. siberu); middorsal colour pattern black,
marked with short or longer line with break in adults and
unmarked in juveniles (vs. black, unmarked); yellow to orange
(vs. usually orange), round or long (vs. round), dense (vs.
sparse) spots on dorsal side of limbs; throat grey to brown
with light spots in life (vs. light grey); abdomen grey to
brown with light reticulation in life (vs. light grey); ventral
skin of throat, chest, abdomen, limbs grey to brown, with
white spots on throat, chest, and sometimes to abdomen (vs.
light grey, usually without spots or reticulation); mean of tibia
length 23.2 mm (vs. 20.9 mm). Morphological comparison
showing dorsal, ventral, palmar and plantar view of these
two species is provided in Fig. 6.
DISCUSSION
In an attempt to infer the phylogeny and revisit the
systematics and biogeography of ranid frogs, Oliver et al.
(2015) elevated numerous phylogenetically distinct sub
genera (including Pulchrana) to genera. The constituents of
Pulchrana previously had been referred to as Hylarana. The
genus Pulchrana, as currently known, is distributed across
Sundaland, and comprises 16 species (Frost, 2018), including
11 species recognised within the P. signata Complex,
namely, P. banjarana, P. centropeninsularis, P. debussyi
(by implication; see Oliver et al., 2015), P. grandocula, P.
mangyanum, P. moellendorf, P. picturata, P. siberu, P.
signata, and P. similis (Brown & Guttman, 2002; Chan et al.,
2014), and the recently described P. guttmani (Brown, 2015).
Both Brown & Siler (2013) and Chan et al. (2014) reported
that Pulchrana siberu and P. centropeninsularis formed a
distinct clade, separate from the remaining species of the P.
signata Complex. Although P. fantastica, new species, comes
from the type locality of an enigmatic congener, P. debussyi
(Van Kampen, 1910), a species with no available genetic
data (Chan et al., 2014), we support Chan et al.’s (2014)
conclusion that P. debussyi is not allied to the P. signata
Complex. Chan et al. (2014) considered morphological

296
Arin et al.: New Pulchrana from Sumatra
Fig. 6. Dorsal (a), ventral (b), palmar (c), and plantar (d) views of Pulchrana fantastica, new species (MZB.Amph.28891, male, holotype,
Aceh); (e–h) P. centropeninsularis (MZB.Amph.28767, male, Jambi); (i–l) P. siberu (BMNH 1979.306, male, holotype, Siberut). Scale
bar = 10 mm. Photo by G. Cahyadi (a–h); U. Arin (i–l).

297
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characters used in the original description P. debussyi and
suggested this species was a probable synonym of Amnirana
nicobariensis.
Our results corroborate Chan et al. (2014) in that, (1)
Pulchrana fantastica, new species, was recovered as the
sister taxon to P. siberu + P. centropeninsularis, and (2)
the clade comprising these species was distinct from the
remainder of the P. signata Complex (Fig. 1). However, our
discovery of P. centropeninsularis on the island of Sumatra
runs contrary to the suggestion by Chan et al. (2014) that
P. centropeninsularis was endemic to the Malay Peninsula.
The record of P. centropeninsularis on Sumatra furthermore
supports the possibility that the shared most recent common
ancestor of P. siberu, P. centropeninsularis, and P. fantastica,
new species, probably originated on Sumatra. Under such a
scenario, P. centropeninsularis may have dispersed to the
Malay Peninsula across the Strait of Malacca. Similarly,
P. siberu may have dispersed to Siberut, Sipora, and Pagai
Selatan across the Strait of Mentawai possibly during the
Pleistocene. At this time period, sea levels were ~120 m lower
and the distance between these landmasses was considerably
narrower (Geyh et al., 1979; Voris, 2000; Chan et al., 2014).
Considering that the genus Pulchrana comprises lineages that
stem from an ancient process of diversication dating back
to the late Eocene (Chan & Brown, 2017), we predict that
several other deeply divergent species probably remain to be
discovered, particularly in the clade containing P. picturata
(Brown & Siler, 2013). This prediction can likely be applied
to more taxa on the large, topographically heterogeneous
island of Sumatra, which remains inadequately sampled. We
suspect that even though the cumulative total of Sumatra’s
amphibians continues to increases every year (Stuart et al.,
2006; Inger et al., 2009; Teynie et al., 2010), its anuran
amphibian diversity will likely remain underestimated for
some time.
Given that the understanding of global amphibian decline is
at a critical stage (Stuart et al., 2004; Whittaker et al., 2013),
comprehensive amphibian surveys are essential to assess the
true diversity of anurans on the island. The documentation
of frog distribution is also of paramount importance for the
design of informed conservation priorities (Rowley et al.,
2010). The IUCN (2017) estimated that 2,067 species of the
globally known 6,533 taxa were threatened, and it is indeed
troubling that almost 2% of these threatened species occur
in Indonesia. Because thorough information concerning the
status of most Indonesian amphibians is lacking, the actual
number of threatened species likely is much higher. The loss
of primary forest (resulting from deforestation and habitat
degradation) is currently the foremost threat for Southeast
Asian amphibians (Rowley et al., 2010). Indonesia has
experienced an unprecedented loss of primary forest, and
between 2000 and 2012, forest loss was at an estimated rate
of 47,600 ha/yr. Within this same period, a staggering 2,857
kha of primary forest loss was recorded in Sumatra, of which
1,205 kha was lowland forest (Margono et al., 2014). This is
a matter of severe concern for the species considered here,
given that Pulchrana siberu, P. centropeninsularis, and P.
fantastica, new species, all exclusively depend on lowland
forests for survival.
ACKNOWLEDGEMENTS
This study was funded by Deutsche Forschungsgemeinschaft
(DFG) under Ha2323/12-1 and stipend was granted for UA
by Deutscher Akademischer Austauschdienst–Indonesian
German Scholarship Program (DAAD–IGSP; 91548731).
AJ works at PT. REKI-Hutan Harapan was funded by DFG
(GL665/4-1) and conducted under RISTEK permit 436/
SIP/FRP/SM/XI/2012 and 81/EXT/SIP/FRP/SM/IX/2013.
Research and collecting permit for UA and GC was issued
by Direktorat Jendral Perlindungan Hutan dan Konservasi
Alam (S.49/KKH-2/2014, SI.10/Set-3/2014). The authors
thank the School of Life Sciences and Technology-Institute
of Technology Bandung, Indonesian Science Institute,
Minister of Research and Technology, Director General
KKH–PHKA as well as Balai Besar Taman Nasional Gunung
Leuser (BBTNGL), Balai Konservasi Sumber Daya Alam
(BKSDA) Aceh, PT. REKI-Hutan Harapan Jambi, Frankfurt
Zoological Society, and BKSDA Jambi which made this
research possible. Authors are also very grateful with Lukman
Shalahuddin and Dadit Herdikiagung, Novari Fajria, Amir
Hamidy, Agus Yasin, Yoghi Budianto, Zainal, Kamarudin,
Carmidi, Hajidin, Zamrin, Agusman, Aidil, Zainudin, Rikha,
Sumarto, Darlizon, Muhardi, Samin, Hasbalah, Alan, Mistar
Kamsi, Andy Arissaputra, Kevin, Musadat and many other
people for all support during eld work in Sumatra. We
would also like to thank Annamarie Vogt, Dimitrij Trovinov,
Katharina Gebauer, Lisa Gottschlich for their support during
lab work. Finally we thank Chan Kin Onn, Rafe Brown,
and Ben Evans for their kind support, Jim McGuire and
Carol Spencer for tissue samples from Mentawai Islands
and Jeffrey Streicher for access to the type specimens of
Pulchrana siberu. Indraneil Das and two other anonymous
reviewers for reviewing this paper.
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APPENDIX
Material examined. Pulchrana centropeninsularis (3). Indonesia—Jambi Province: Hutan Harapan, MZB.Amph.28765–67.
Pulchrana fantastica, new species (18). Indonesia —Aceh Province: Mane, MZB.Amph.28895; Taman Buru Linge Isaq,
MZB.Amph.28889–94, MZB.Amph.28896–98, MZB.Amph.28943–45, MZB.Amph.28947–48; Taman Nasional Gunung
Leuser, MZB.Amph.28946. Sumatera Utara Province: Bandar Baru, MZB.Amph.13011, MZB.Amph.13015.