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A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a Phylogeny of Sundaland Toad Genera


Abstract and Figures

We describe a new genus and two new species of toads from the Sumatran volcanoes Gunung Sorikmarapi and G. Kunyit, in the provinces of Sumatera Utara and Jambi, respectively. The new taxa can be distinguished from other genera, and each other, based on genetic differentiation, morphology, and advertisement call structure. We employ both nuclear and mitochondrial data to provide a phylogenetic hypothesis of relationships for the bufonid genera of the Sunda Shelf. While broadly corroborating previous studies, our results also shed light on the phylogenetic position of Pseudobufo. The new genus, Duttaphrynus, and Pseudobufo are basal to other Sunda Shelf genera in our phylogenies.
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A New Genus and Two New Species of Arboreal Toads from the Highlands of
Sumatra with a Phylogeny of Sundaland Toad Genera
Author(s): Utpal Smart, Goutam C. Sarker, Umilaela Arifin, Michael B. Harvey, Irvan Sidik, Amir
Hamidy, Nia Kurniawan, and Eric N. Smith
Source: Herpetologica, 73(1):63-75.
Published By: The Herpetologists' League
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Herpetologica, 73(1), 2017, 63–75
Ó2017 by The Herpetologists’ League, Inc.
A New Genus and Two New Species of Arboreal Toads from the Highlands of Sumatra with a
Phylogeny of Sundaland Toad Genera
Amphibian and Reptile Diversity Research Center and Department of Biology, University of Texas at Arlington, Arlington, TX 76019, USA
Centrum f ¨ur Naturkunde - Zoologisches Museum, Universit¨
at Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
Department of Biological Sciences, Broward College, 3501 S.W. Davie Road, Davie, FL 33314, USA
Laboratory of Herpetology, Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences–LIPI, Widyasatwa Loka Jl.
Raya Jakarta Bogor km 46, Cibinong, West Java, Indonesia
Department of Biology, Universitas Brawijaya, Jl. Veteran, Malang, East Java, Indonesia
ABSTRACT: We describe a new genus and two new species of toads from the Sumatran volcanoes Gunung Sorikmarapi and G. Kunyit, in the
provinces of Sumatera Utara and Jambi, respectively. The new taxa can be distinguished from other genera, and each other, based on genetic
differentiation, morphology, and advertisement call structure. We employ both nuclear and mitochondrial data to provide a phylogenetic
hypothesis of relationships for the bufonid genera of the Sunda Shelf. While broadly corroborating previous studies, our results also shed light on
the phylogenetic position of Pseudobufo. The new genus, Duttaphrynus, and Pseudobufo are basal to other Sunda Shelf genera in our phylogenies.
Key words: Barisan Range; Bufonidae; Molecular phylogenetics; Sunda Shelf; Taxonomy
BETWEEN June 2013 and February 2014, we carried out an
inventory of the highland herpetofauna of Sumatra. One trip,
in January 2014, revealed a very peculiar species of
unidentified toads from the slopes of Gunung Sorikmarapi,
a densely vegetated stratovolcano situated in Batang Gadis
National Park, in the Mandailing Natal Kabupaten of the
Sumatera Utara Province. We found the first specimen
inside a small subterranean hollow, crouching motionless on
a rock wall, ~2 m below the ground. Further searching
inside caves in the area did not reveal more individuals.
Nonetheless, we found three more specimens at nearby
localities the following day. While comparing these speci-
mens to others collected from the previous year, we came
across another individual that was collected at Gunung
Kunyit of the Barisan Range in Jambi Province, Sumatra in
June 2013, which exhibited close morphological affinities to
the peculiar specimens from Batang Gadis National Park.
Here we appraise these specimens within the framework
of a lineage-based, Unified Species Concept (de Queiroz
2005). Using morphological diagnosability, phylogenetic
relationships, genetic divergence, and patterns of advertise-
ment calls for assessing the evolutionary independence of
this lineage, we provide several lines of evidence to validate
the recognition of this lineage as a new genus and its
constituent taxa as two new species. We also take this
opportunity to clarify the genus-level phylogeny of the toads
found on the Sunda Shelf.
Specimens used in this work were collected during
fieldwork in Sumatra and Java between May 2013 and
August 2015. We photographed specimens when alive and
then again after they were fixed according to Institutional
Animal Care and Use Committee protocols. We took dorsal,
ventral, and lateral photographs of the specimens post
mortem and with a scale for size reference. We preserved
specimens in a 1:9 dilution of stock formalin in water until
they could be transferred to 70% ethanol. Finally, we
deposited specimens at the Museum Zoologicum Bogoriense
(MZB), Cibinong, Indonesia, and the Amphibian and Reptile
Diversity Research Center of the University of Texas at
Arlington (UTA), Texas, USA.
We used morphological terminology primarily based on
Matsui (1984), while also using terminology of Duellman
(2001) and Kok and Kalamandeen (2008). With a digital
caliper (3 mm) or ocular micrometer (,3 mm) to the
nearest 0.1 mm, we measured: snout–vent length (SVL);
head length; head width (width of the head measured at the
angles of the jaw, excluding warts at the jaw); snout length
(from the anterior ocular angle to the tip of the snout);
intercanthal distance (distance between anterior edges of
canthus); internarial distance (distance between proximal
ends of nares); interorbital distance (the minimal distance
between the upper eyelids); eye length (distance between
the anterior and posterior adjunction of upper and lower
eyelids); eye–nares length (distance from the proximal
junction of upper and lower eyelids and proximal end of
nares); nares–snout length (distance between the proximal
end of nares to the tip of the snout); tympanum width
(horizontal); forearm length (from tip of the elbow to the
proximal margin of outer metatarsal tubercle); hand length
(from the distal margin of the metatarsal tubercle to the tip
of the third finger); femur length, tibia length (the greatest
length of tibia by positioning the hind limb in a Z pattern);
tarsus length (distance of tibio-tarsal joint to the proximal
end of outer metatarsal tubercle); foot length (from the distal
margin of the outer metatarsal tubercle to the tip of Toe IV);
and finger pad width (greatest width of finger pad). We used
the webbing formulae of Myers and Duellman (1982). We
used digital images to describe color in life; these images are
deposited at the University of Texas at Arlington (UTA)
digital image collection. We followed the museum acronyms
available in Sabaj Perez (2014).
Molecular Phylogenetic Analyses
Taxon sampling and DNA sequencing.—We included
21 individual operational taxonomic units of 13 distinct toad
genera in this study (Table 1). Apart from the three
undescribed specimens, this data matrix also included
sequences from type species of all genera of toads that
occur in the Sunda Shelf, except for Pelophryne whose type
species, P. albotaeniata (Barbour 1938), lacks sequence data.
In our matrix, Pelophryne was thus represented by P.
brevipes and P. signata. To minimize the effect of long-
branch attraction, and to help stabilize the phylogeny (see
Bergsten 2005), we also included one species of Sumatran
toad from each genus alongside the types where possible.
Pedostibes hosii was recently made the type species for the
new genus Rentapia (Chan et al. 2016), but previously
belonged to a genus that encompassed an Indian endemic
(Pedostibes tuberculosus, type species of Pedostibes) and four
Southeast Asian arboreal toads (Frost et al. 2006). We
decided to incorporate Pedostibes tuberculosus in our study,
to contextualize our phylogeny in the light of this recent
taxonomic development. To minimize missing data, we chose
not to include R. rugosus in our analysis because only the
small ribosomal subunits 12S and 16S are currently available
for this species on GenBank. Our sampling of Bufonidae
thus incorporated species of the following genera: Ansonia
(A. penangensis type species and A. leptopus); Duttaphrynus
(D. melanostictus type species); Ingerophrynus (I. biporcatus
type species and I. divergens); Leptophryne (L. cruentata
type species and L. borbonica); Pedostibes (P. tuberculosus
type species); Rentapia (R. hosii type species); Pelophryne
(P. signata and P. brevipes); Sabahphrynus (S. maculatus
type species); the monotypic genus Pseudobufo (P. subasper
type species); and two undescribed specimens from Gunung
Sorikmarapi and one undescribed specimen from Gunung
Kunyit. We also included American Toads (Anaxyrus
americanus) and Canyon Treefrogs (Dryophytes [Hyla]
arenicolor) as outgroup taxa (Duellman et al. 2016). Here,
we provide new sequences for 11 individuals; the remaining
10 were published by Van Bocxlaer et al. (2009) and
obtained from GenBank.
Prior to preservation of specimens, we collected either
muscle or skin tissue and stored it in 1.5 mL of cell lysis
buffer solution (0.5 M Tris/0.25% EDTA/2.5% SDS, pH ¼
8.2) or in 95% ethanol. We isolated Genomic DNA using a
Qiagen DNeasy kit (Qiagen, Valencia, CA, USA) or AMPure
XP beads following the Agencourt protocol (Beckman
Coulter Co., Fort Collins, CO, USA) after Rohland and
Reich (2012). We sequenced 2 mitochondrial (12S and 16S)
and 2 nuclear genes (NCX1 and CXCR4), slightly modifying
the primers and thermocycle protocols provided by Van
Bocxlaer et al. (2009). We used these loci because they have
been screened by previous studies on bufonids and have
been established as informative (Biju et al. 2009; Van
Bocxlaer et al. 2009, 2010). We used the GoTaqtGreen
Master Mix, 2X (Promega Corporation, Madison, WI, USA)
for all amplification reactions on a GeneAmptPCR System
9700 (Applied Biosystems, Foster City, CA, USA). We
checked all successful polymerase chain reaction products
visually in agarose gels and purified them using AMPure XP
beads following the Agencourt protocol. The Genomic Core
Facility at the University of Texas at Arlington completed the
sequencing reactions with an ABI PRISM 3100xl Genetic
Analyzer (Applied Biosystems).
TABLE 1.—Specimens used in a molecular analysis of bufonid anurans from the Sunda Shelf, Malaysia. If museum numbers are unavailable, we provide
the field number (SDB) or the tissue collection number (VUB). New sequences are indicated in bold.
Species Locality Museum number
Genbank accession number
12S 16S NCX1 CXCR4
Dryophytes (Hyla)arenicolor USA; Arizona TNHC 61118 (VUB 1052) FJ882776 FJ882776 EF107241 AY364190
Anaxyrus americanus USA; Mississippi CAS 207258 FJ882827 FJ882827 FJ882676 FJ882730
Ansonia leptopus Malaysia; Borneo VUB 0632 FJ882795 FJ882795
Ansonia penangensis Malaysia; Penang KUHE UNL1 AB435262 AB435262
Duttaphrynus melanostictus Indonesia; Java UTA 53737 KX192078 KX192086 KX19211
Ingerophrynus biporcatus Indonesia; Sumatra UTA 53730 KX192079 KX192090 KX19212
Ingerophrynus divergens Malaysia; Borneo VUB 0602 FJ882802 FJ882802 FJ882648 FJ882701
Leptophryne cruentata Indonesia; Java UTA 62522 KX192075 KX192087 KX19207 KX192108
Leptophryne borbonica Indonesia; Sumatra UTA 62486 KX192076 KX192095 KX19208
Pedostibes tuberculosus India; Western Ghats SDB 4691 FJ882793 FJ882793 FJ882640 FJ882693
Pelophryne brevipes Indonesia; Sumatra UTA 63762 KX192080 KX192088 KX19210
Pelophryne signata Malaysia; Borneo VUB 0583 FJ882801 FJ882801 FJ882646 FJ882699
Phrynoidis juxtasper Malaysia; Borneo VUB 0649 FJ882805 FJ882805 FJ882656 FJ882710
Phrynoidis asper Indonesia; Java UTA 53719 KX192077 KX192089 KX19209
Pseudobufo subasper Indonesia; Sumatra UTA 63763 KX192083 KX192096 KX19216
Pseudobufo subasper Indonesia; Sumatra UTA 63764 KX192084 KX192093 KX19217
Rentapia hosii Malaysia; Borneo VUB 0661 FJ882804 FJ882804 EF107223 EF107449
Sabahphrynus maculatus Malaysia; Sabah BORNEENSIS 08425 AB331718 AB331718
Sigalegalephrynus mandailinguensis
gen. nov., sp. nov. Indonesia; Sumatra UTA 63562 KX192082 KX192092 KX19215 KX192110
Sigalegalephrynus mandailinguensis
gen. nov., sp. nov. Indonesia; Sumatra MZB 25736 KX192081 KX192094 KX19214 KX192109
Sigalegalephrynus minangkabauensis
gen. nov., sp. nov. Indonesia; Sumatra MZB 25738 KX192085 KX192091 KX19213
64 Herpetologica 73(1), 2017
Sequence alignment and phylogeny inference.—We
assembled and cleaned the raw gene-fragment sequences
using Sequencher (v4.8; Gene Codes, Ann Arbor, MI, USA).
Prior to alignment, we used the Gblocks Server (v0.91b;
Castresana 2000; Talavera and Castresana 2007) to eliminate
poorly aligned and hyper-variable regions present in the
ribosomal subunits. We then obtained the alignment of
sequences using Clustal W (Larkin et al. 2007) implemented
in MEGA (v5.1; Tamura et al. 2011). We translated the
sequences of the protein coding genes to amino acid
sequences, to verify the absence of stop codons and proper
alignment, and where necessary, we edited them by eye for
accuracy. We did not detect any internal stop codons and we
deposited the new sequences in GenBank (accession
numbers in Table 1).
We partitioned the concatenated data set by gene and
codon position and used Partition-Finder (v1.1.0; Lanfear et
al. 2012) to determine the best partitioning schemes and
models for each partitioned subset. Partition-Finder simul-
taneously determined the partition schemes based on
Bayesian Information Criterion using the ‘‘ greedy’’ search
algorithm. The best partitioning scheme had seven parti-
tions: 12S and 16S, GTRþIþC; CXCR4 codon 3, HKYþI;
CXCR4 codon 1, HKYþG; CXCR4 codon 2, GTR; NCX1
codon 2, HKYþI; NCX1 codon 3, GTRþC; and NCX1 codon
1, HKYþIþC.
To examine the phylogenetic position of the undescribed
species in relation to Sundaland toads, we used maximum
likelihood (ML) and Bayesian inference (BI) methods. We
conducted ML analysis employing the rapid bootstrapping
algorithm using the program RAxML v8.00 (Stamatakis
2014) on the CIPRES Science Gateway server (v3.2; Miller
et al. 2010). Because the 25 discrete rate categories are said
to better approximate invariant sites (Stamatakis 2006), we
used the model GTRþG instead of GTRþIþG for the ML
analysis. Nodal support for ML was provided by boot-
strapping (BS; 1000 pseudoreplicates), with BS values 0.70
considered strong support (Hillis and Bull 1993).
We conducted Bayesian Markov chain Monte Carlo
(MCMC) phylogenetic analyses on a partitioned alignment
using MrBayes (v3.3; Ronquist and Huelsenbeck 2003). We
initiated two simultaneous runs of four MCMC analyses,
consisting of one cold and three incrementally heated chains,
with random trees for 5 310
total generations (sampling
every 500 generations). We set the burn-in to the default
values of 25%, hence discarding the initial 2500 generations.
To examine stationarity, we used trace plots and ESS values
(.200) on TRACER v1.5 (Rambaut and Drummond 2009).
We constructed a 50% majority-rule consensus tree with
estimates of Bayesian support using the remaining sampled
trees and posterior probabilities (PP), wherein PP values
0.95 were considered strong support (Alfaro et al. 2003;
Huelsenbeck and Rannala 2004; Mulcahy et al. 2011). We
used the graphical viewer Figtree (Rambaut 2007) to edit the
resulting output of RAxML and MrBayes analyses.
Advertisement Call Analysis
Elijah Wostl recorded the vocalization of an uncaptured
individual from near Lake Saba Begu on 26 January 2014.
He recorded a single call at 2156 h using a Zoom H4n Handy
Recorder at a sampling rate of 44.1 kHz. The ambient
temperature at the time of the recording was 18.78C. To
remove background noise, we first filtered the recording
using the free sound-editing software Audacity (v2.0.3;
Audacity Team 2014).
We analyzed the temporal and spectral characteristics of
the recorded vocalizations using the sound synthesis and
analysis package Seewave (v1.7.3; Sueur et al. 2008)
implemented in RStudio (v0.98.1062; R Core Team 2013).
We measured four traditional call characters known to be
important in communication, including dominant frequency,
call duration, pulse duration, and pulse rate (Cocroft and
Ryan 1995). We estimated dominant frequency in Seewave
using a fast Fourier transformation (Hanning window length
¼100 samples; 85% overlap between successive windows).
We obtained two-dimensional spectrograms using the
function spectro. We measured temporal properties using
the function timer, with a 2% amplitude threshold for signal
detection for pulse duration and pulse rate.
Phylogenetic Analyses
Our final data matrix consisted of 1537 base pairs and was
94% complete, with only two species lacking both nuclear
loci (Ansonia penangensis and Sabahphrynus maculatus) and
one individual of the undescribed species from Gunung
Sorikmarapi for which we were unable to amplify the
CXCR4 exon. Our ML analyses produced a single tree (lnL
¼6110.25; Fig. 1) that is almost identical to our Bayesian
consensus phylogram. Most basal nodes are poorly support-
ed; hence, the relationships among the Sundaland toad
genera remain essentially unresolved. Each genus is
recovered as monophyletic with high support. The group
containing the undescribed individuals from Sumatra is
nested within the Sundaland toads and forms a well-
supported clade (BS ¼100; PP ¼1) with the two specimens
from Gunung Sorikmarapi being sister to a more divergent
specimen from Gunung Kunyit (uncorrected ‘‘p’’ distance ¼
4.2%; Table 2; BS ¼100; PP ¼1). We recovered a strongly
supported sister relationship between Pedostibes tuber-
culosus and Duttaphrynus melanostictus (BS ¼80; PP ¼
98), and together these two genera form a sister relationship
with all Sundaland bufonids (BS ¼14; PP ¼0.96). We also
recovered a moderately strong sister relationship between
Rentapia hosii and frogs of the genus Phrynoidis (BS ¼52;
PP ¼96), again in concordance with previous molecular
phylogenies (Matsui et al. 2007; Van Bocxlaer et al. 2009,
2010; Pyron and Wiens 2011; Chan et al. 2016). Our analyses
recovered the genus Ansonia as sister to a clade containing
Pelophryne and Ingerophrynus with modest support in both
analyses (BS ¼25; PP ¼0.77). Ansonia and Pelophryne have
been recovered as sister taxa before (Van Bocxlaer et al.
2009; Pyron and Wiens 2011) but to the exclusion of
Ingerophrynus. Our analyses did not confirm the sister
relationship between Ingerophrynus and Sabahphrynus
(contra Van Bocxlaer et al. 2009; Pyron and Wiens 2011),
but instead indicated weak support for the latter as sister to
Leptophryne (BS ¼14; PP ¼0.47). The monotypic genus
Pseudobufo aligned with D. melanostictus and P. tuber-
culosus in our ML tree (BS ¼20); whereas in our BI tree, it
was recovered as sister to all Sundaland toads (PP ¼0.44).
Despite poor support, both analyses assign a basal position to
Pseudobufo relative to all other Sundaland bufonid genera.
The results of our phylogenetic analyses, as well as the call
pattern data, indicate that the two undescribed species of
toads form a distinct lineage among Southeast Asian
bufonids. Therefore, we propose to establish a new genus
for these two species.
Sigalegalephrynus gen. nov.
Type species.Sigalegalephrynus mandailinguensis by
present designation.
Diagnosis and comparisons.—The genus can be
diagnosed based on the following ecological attributes and
morphology: medium-sized (,40 mm SVL) member of
Bufonidae, with gangly limbs, and arboreal and troglodytic
habits. The morphologically comparable genus Ansonia
(.40 mm SVL), commonly called Stream toads, is typically
found on low vegetation near watercourses. The generally
diminutive members of Pelophryne (,40 mm SVL) and
Leptophryne (.40 mm SVL) occur on shrubs. The only
other genus of toads that has true arboreal habits is Rentapia,
whose members typically tend to be stockier (.70 mm SVL)
than Sigalegalephrynus. The genus Phrynoidis is represent-
ed by two large (.70 mm SVL) semiaquatic toads, usually
found on rocks along streams and rivers; the members of the
Duttaphrynus (.40 mm SVL) and Ingerophrynus (.40 mm
SVL) display terrestrial or somewhat riparian habits. The
monotypic genus Pseudobufo is represented by a large (.75
mm SVL) and aquatic species with completely webbed feet
that inhabits the peat swamps of the Malay Peninsula,
Borneo, and Eastern Sumatra.
Like all other Sundaland toad genera (characters in
parentheses), the new genus possesses a visible or slightly
visible tympanum. Sigalegalephrynus most closely resembles
Ansonia; however, the former lacks mandibular spines
(mandibular spines present) and possesses combined femur
and tibia lengths smaller than its SVL (SVL ,femur and
tibia lengths). Unlike Pelophryne, in Sigalegalephrynus
Finger I projects beyond the webbing by two phalanges
(reduced Finger I, with one or no phalanges projecting
beyond webbing), and males possess nuptial excrescences
with well-keratinized spicules (poorly spiculated with only
slight keratinization or not keratinized at all). Unlike
Rentapia, Sigalegalephrynus lacks paratoid glands (paratoids
prominent). The new genus can be told apart from
Ingerophrynus by the lack of well-defined parallel crests
between the eyes (parallel crests prominent). Unlike
FIG. 1.—Phylogeny of Sundaland toads, based on 1537 base pairs of a combined mitochondrial and nuclear data set, depicted as a maximum-likelihood
consensus tree with Anaxyrus americanus as a proximate outgroup (the nonbufonid outgroup Dryophytes [Hyla]arenicolor has been trimmed out).
TABLE 2.—Pairwise genetic distances (uncorrected p) observed in the
sequence of the mitochondrial 16S ribosomal subunit gene between two
new species of puppet toads, and between arboreal toad genera of
Pairwise comparison % difference
S. mandailinguensis to S. minangkabauensis 4.2
Sigalegalephrynus to Pelophryne 9–9.55
Sigalegalephrynus to Leptophryne 7.55–8.75
Sigalegalephrynus to Ansonia 8.6
Sigalegalephrynus to Ingerophrynus 6.55–8.25
Sigalegalephrynus to Rentapia 8.6
Sigalegalephrynus to Pedostibes 6.1
Sigalegalephrynus to Phrynoidis 6.75–6.95
Sigalegalephrynus to Sabahphrynus 8.2
Sigalegalephrynus to Duttaphrynus 6.85
Sigalegalephrynus to Pseudobufo 8.0
Pelophryne to Leptophryne 8–10.85
Pelophryne to Ansonia 10.1
Pelophryne to Rentapia 9.9
Pelophryne to Sabahphrynus 10.5
Ansonia to Rentapia 7.0
Ansonia to Sabahphrynus 9.4
Sabahphrynus to Rentapia 9.4
66 Herpetologica 73(1), 2017
Leptophryne, Sigalegalephrynus lacks enlarged tubercles at
the base of each toe, between at the articulation of the first
phalanx and metacarpus (large tubercles present). Addition-
ally, males of L. cruentata are unique among Southeast Asian
toads in having nuptial excrescences that are white and
swollen on the first and second fingers; males of Sigalega-
lephrynus (and all other genera) have nuptial excrescences
with no white and swollen tissue. Unlike Phrynoidis,
Sigalegalephrynus has slender limbs (limbs robust) and toes
that are less than half webbed (toes fully webbed, with the
exception of the fourth). Unlike Pseudobufo—the only
Sundaland toad with fully webbed toes—Sigalegalephrynus
has toes that are less than half webbed.
Males of Sigalegalephrynus can be distinguished from all
other toads in the region by the presence of an elongate
inner metacarpal-thenar tubercle, which is as distinct and
large as the outer metacarpal tubercle, and is located
medially (Fig. 2). Males of Leptophryne cruentata have an
elongate and medially located inner metacarpal tubercle, but
this is less distinct and noticeably smaller than the outer
metacarpal tubercle, whereas Rentapia and Pelophryne lack
the inner metacarpal tubercle altogether. Fingertips three
and four of the new genus are truncated, reflecting
arboreality as in Pelophryne, Sabahphrynus, Rentapia, and
some species of Ansonia.
Etymology.—The generic name is derived from the
name given by the indigenous Batak people of the Toba
region in Sumatera Utara to life-sized wooden puppets
called Sigale Gale. These puppets are used during the
papurpur sepata funerary festivals to placate the spirits of
the dead who have left no children behind. The suffix is
derived from the masculine and Latinized Greek noun for
toad, phrynos. The new genus, with a relatively large size
compared with most arboreal toads in the region, lanky
hands, and a wood-brown complexion, is evocative of the
Sigale Gale.
Common name.—Puppet Toads.
Content.—Our phylogenetic analyses indicate the pres-
ence of two species within the new genus: S. mandailin-
guensis Smart et al.; and S. minangkabauensis Smart et al.
Sigalegalephrynus mandailinguensis sp. nov.
Holotype.—Museum Zoologicum Bogoriense Amphibian
Collection, MZB 25736 (field number ENS 15697; Fig. 3),
an adult male from above the village of Sibanggor Tonga on
the northeast slope of Gunung Sorikmarapi, Kecamatan
Panyabungan Selatan, Kabupaten Mandailing Natal, Provinsi
Sumatera Utara, Indonesia, 0.701648N, 99.552628E, 1383 m
(in all cases, datum ¼WGS84), collected by U. Smart, I.
Sidik, and E.N. Smith on 25 January 2014.
Paratypes (3).—UTA 63561 (field number ENS 16709),
adult male from trail between the Tano Bato to Sapo Tinjak
road and Lake Saba Begu, Batang Gadis National Park,
Kecamatan Batang Natal, Kabupaten Mandailing Natal,
Provinsi Sumatera Utara, Sumatra, Indonesia 0.708458N,
99.518998E at 1299 m. UTA 63562 (field number ENS
16936) and MZB 25737 (field number ENS 16937), adult
males from the same locality as previous paratype but at
1297 m 0.708668N, 99.519538E. All paratypes collected by
U. Smart, S. Handayani, and I. Sidik on 26 January 2014.
Diagnosis.—The following combination of characters is
unique to Sigalegalephrynus mandailinguensis: (1) This
medium-sized (males 30.6–38.0 mm SVL) slender toad lacks
parotoid glands. (2) The tympanum is visible. (3) The nares
are closer to the tip of the snout than to the eye. (4) The
fingertips are truncated and expanded. (5) The toe tips are
truncated but not expanded. (6) The webbing is rudimentary
in the hands and moderate in the feet. (7) The dorsal
coloration consists of white and brown with a thin stripe
extending from the tip of the snout to the vent at midline. (8)
A dark band above orbits is joined medially, and extends as
an interrupted medial track to the sacrum. (9) Alternate
black–dark brown and white marks on the upper lip. (10)
The flanks have a stroke of brown extending from the orbit to
the inguinal area. (11) The dorsal surface is moderately
tuberculate. (12) The surface of the abdomen is uniformly
tuberculate, with small, smooth, and round tubercles.
Description of holotype and variation.Holotype
(adult male) followed by variation of three adult male
paratypes in parentheses (UTA 63561; UTA 63562; MZB
25737). The specimen has SVL of 38.0 mm (30.6, 32.29,
32.75); head length 11.34 mm (9.5, 9.73, 10.19); head width
11.3 mm (9.65, 10.27, 10.22); snout length 5.0 mm (4.0, 3.6,
3.5); internarial distance 3.0 mm (2.5, 2.68, 2.75); eye length
FIG. 2.—Palmar (upper) and plantar (lower) surfaces of Sigalegaleph-
rynus mandailinguensis (A; MZB 25736) and S. minangkabauensis (B; MZB
3.3 mm (2.8, 3.1, 3); interocular distance 4 mm (3.5, 3.5, 3.7);
intercanthal distance 5.7 mm (4.4, 5.0, 5.5); tympanum width
2.1 mm (1.5, 1.8, 1.8); hand length 11.2 mm (9.4, 9.1, 9.8);
forearm length 11.5 mm (10, 9.6, 10.3); femur length 15.85
mm (13.46, 12.88, 13.38); tibia length 17.0 mm (13.86, 13.1,
14.02); tarsus length 9.0 mm (7.3, 7.2, 7.5); foot length 16.37
mm (13.58, 13.04, 13.68); width of fingertip pads for Finger I
0.10 mm (0.09, 0.08, 0.09), Finger II 0.13 mm (0.11, 0.10,
0.11), Finger III 0.14 mm (0.11, 0.11, 0.13), Finger IV 0.14
mm (0.10, 0.11, 0.13); width of toe pads for Toe I 0.09 mm
(0.07, 0.07, 0.07), Toe II 0.09 mm (0.08, 0.08, 0.08), Toe III
0.11 mm (0.08, 0.08, 0.09), Toe IV 0.12 mm (0.09, 0.09,
0.10), Toe V 0.12 mm ( 0.09, 0.09, 0.11).
Body slender; head almost as long as wide; head length
30% (31%, 30%, 31%) of SVL; head width 30.0% (32%, 32%,
31%) of SVL; snout length 13% (13%, 11%, 11%) of SVL;
canthus rostralis concave; loreal area slightly tuberculate and
concave; eye length 8.7% of SVL; pupil horizontal; snout
slightly sloping back toward mouth; snout mucronate and
with prominent median keel, in dorsal view; tympanum
distinct, with moderately developed supratympanic fold;
interorbital space flat; cranial crests absent; no teeth in jaws;
tongue tip oval-shaped and longer than wide; skin of dorsal
FIG. 3.—(A) Lateral, (B) dorsal, and (C) ventral view of the adult male holotype of Sigalegalephrynus mandailinguensis from Gunung Sorikmarapi,
Sumatera Utara Province, Sumatra (MZB 25736). A color version of this figure is available online.
68 Herpetologica 73(1), 2017
surfaces rough to finely shagreen with few large, scattered
tubercles; tubercles small, rounded, and almost without
keratinization; no dorsolateral, paravertebral, or occipital
folds; skin on venter smooth with very fine warts; forearm
length 30.3% (32.7%, 29.7%, 31.5%) of SVL; hand length
29.5% (30.7%, 28.2%, 29.9%) of SVL; relative length of
Finger I ,II ,IV ,III; fingers bearing large, expanded
pads; webbing formula for hand: I1
2IV (I[2–2], [2–2], [1
–2] II[1
], [1
], [1
], [2
–2], [2
]IV); skin of forearm
with tubercles; Finger I with elongate inner metacarpal
tubercle, as large as outer metacarpal tubercle; each finger
with one poorly developed round subarticular tubercle;
nuptial excrescence brownish-dark, glandular, and dorsome-
dially extended with keratinized spicules present at the base
of Finger I; femur length 41.7% (44%, 39.9%, 40.9%) of
SVL; tibia length 44.8% (45.3%, 40.6%, 42.8%) of SVL;
tarsal length 23.7% (23.9%, 22.3%, 22.9%) of SVL; foot
length 43.1% (44.4%, 40.4%, 41.8%) of SVL; relative lengths
of toes I ,II ,III ,V,IV; toes bearing large pads; feet
with moderate webbing, webbing formula for the feet: I0–
V (I[0–1], [1–1
], [0–
]II[1–2], [1–2], [1–2]III[1
–3], [1
], [1–
–2], [2
], [2
]V); heels without
tubercles; inner metatarsal tubercle moderately developed
and elongate; outer metatarsal tubercle absent; one moder-
ate subarticular tubercle present at the base of first phalanx
on each toe; toes without toe pads.
Color in life.—Distinct light brown spot on each side of
lore and distinct whitish spot on upper jaw just below
posterior end of orbit; dorsum brown with light chamois-
brown-colored hourglass marking; distinct darker spots
dorsolateral of sacral and iliac joint; alternate black and
brown bands on dorsal side of limbs; in lateral flanks, skin
with distinct blackish stripe extending from posterior end of
eyes through top of humeral–ulnar joint to inguinal area;
short, black dorsal stripe on each side of pubic junction;
venter yellowish-white maculated with black blotches; chest
and ventral surface of limbs light tea-rose orange with few
sparsely located yellowish dots; tips of fingers and toes tea-
rose orange-colored without dots; iris bright gold with black
Color in preservative.—In preservative the color of the
animal differs slightly from that in life. The dorsum has a
very light hourglass marking. The venter is whitish grey
maculated with dark brown blotches.
Etymology.—The specific epithet is an adjective refer-
ring to the Mandailing Batak ethnic region and Kabupaten
(regency) where the new species was found.
Common name.—Mandailing Puppet Toads.
Distribution and natural history.Sigalegalephrynus
mandailinguensis is known only from rainforests on the
slopes of Gunung Sorikmarapi in southern Sumatera Utara
from 1297 m to 1383 m (Figs. 4 and 5). The holotype was
found at 1830 h inactive on a cave wall, ~2 m from the
entrance and below ground level. The paratypes were active
on vegetation 0.5 m, 2 m, and 3 m (on a liana) above ground
between 1930 and 2115 h. The two individuals higher above
the ground were vocalizing using a series of high-pitched
ticks. The holotype weighed 2.5 g, and the paratypes 1.4, 1.7,
and 1.8 g.
Call.—The advertisement call of Sigalegalephrynus man-
dailinguensis is best described as a rapid succession of
regularly placed, shrill ticks. A single call 17.27 s in length
was recorded at 2156 h. The call begins with infrequent, low-
amplitude ticks, which progressively get louder and attain a
uniform pulse rate during the main calling period that, in our
recording, lasted ~15 s. During this time, the tonal pulses
were organized in a discreet series repeated at a moderately
high rate of 6 times/s. Each individual pulse lasted about 29
64 ms (Fig. 6) intersected by pauses ~12 69 ms long.
These pulses were relatively narrow band (,500 Hz). The
average dominant frequency of pulses was 3.4 60.04 Hz.
Sigalegalephrynus minangkabauensis sp. nov.
Holotype.—Museum Zoologicum Bogoriense Amphibian
Collection, MZB 25738 (field number ENS 16028), an adult
male (Fig. 7) from Gunung Kunyit, Kecamatan Panyabungan
Selatan, Kabupaten Kerinci, Provinsi Jambi, Indonesia,
2.260138S, 101.495128E at 1402 m, collected by E. Wostl,
E.N. Smith, W. Trilaksono, and G. Barraza on 24 June 2013.
Diagnosis and comparison.—The following combina-
tion of characters is unique to Sigalegalephrynus minangka-
bauensis: (1) A small (19.32 mm SVL) and slender toad
without parotoid glands. (2) Fingertips I and II are rounded
and not expanded. (3) Fingertips III and IV are rounded and
expanded. (4) The toe tips are rounded but not expanded. (5)
The webbing is rudimentary in the hands and moderate in
the feet. (6) The dorsum is light greenish-brown with a
middorsal pinstripe extending from the tip of the snout to
the vent. (7) The flanks have a single stroke of dark brown
extending from the posterior end of the orbit to the inguinal
region. (8) The dorsal surface is moderately tuberculate. (9)
The ventral surface is smooth with scattered black spots.
Sigalegalephrynus minangkabauensis can be distin-
guished from S. mandailinguensis (characters in parenthe-
ses) based on the following differences. The tympanum is
barely discernible (tympanum distinct). The finger-pads are
moderately defined (finger-pads prominent). The fingertips
are rounded but not expanded (distinctly spatulate on tips III
to IV). The hands lack subarticular tubercles (distinct
subarticular tubercles under Fingers III and IV). The
webbing of the foot is more extensive, extending to the last
phalanx on Toes I and II (last phalanx free of webbing). The
pads on toes are moderately defined (pads prominent). The
feet lack subarticular tubercles (distinct subarticular tuber-
cles under Toes IV and V). The overall texture is glossy with
fewer tubercles on the dorsum and flanks (overall texture
rugose, body and flanks extensively tuberculate). Taking into
account the uncorrected genetic distance between the two
species of Sigalegalephrynus (Table 2), the aforementioned
comparisons provide adequate diagnostic characters to
warrant S. minangkabauensis as a species distinct from S.
Description of holotype.—The holotype (sex indistin-
guishable) has SVL of 19.32 mm; head length 6.94 mm; head
width 6.57 mm; snout length 2.7 mm; eye length 2.2 mm;
eye–nares length 1.6 mm; distance between nares to tip of
snout 0.6 mm; internarial distance 1.8 mm; intercanthal
distance 3.6 mm; forearm length 6.0 mm; hand length 5.5
mm; femur length 8.5 mm; tibia length 8.71 mm; tarsal
length 5.1 mm; foot length 7.5 mm; width of fingertip pad of
Finger III 1.6 mm; Finger IV 1.4 mm.
Body slender, head little longer than wide; head length
36.1% SVL, head width 34.0% SVL; snout length 14% SVL;
canthus rostralis concave; loreal area without tubercules and
concave; snout truncated and slightly sloping back toward
mouth; snout mucronate and with prominent median keel in
dorsal view; eye length 11.4% SVL; pupil horizontal; upper
eyelid granular; tympanum barely visible, with no supra-
tympanic fold; interorbital space flat; cranial crests absent;
no teeth in jaws; tongue tip oval-shaped, longer than wide;
skin of dorsal surfaces rough to finely shagreen with few
large, scattered tubercles; tubercles small, rounded, and
almost without keratinization; no dorsolateral, paravertebral,
or occipital folds; skin on venter smooth with few fine warts;
forearm length 31.1% SVL; hand length 28.5% SVL; relative
lengths of Fingers I ,II ,IV ,III; fingers bearing
expanded pads; webbing of hands moderate: between
Fingers I and II reaches distal and proximal subarticular
tubercles, respectively; between Fingers II and III reaches
proximal subarticular tubercles; between Fingers III and IV
reaches proximal subarticular tubercles; webbing formula for
the hand I1
–2IV; elongate inner meta-
carpal tubercle below Finger I as large as outer metarcarpal
tubercle; lower arm with indistinct tubercles; fingers without
expanded pads; femur length 44.0% SVL; tibia length 45.0%
SVL; tarsal length 26.3% SVL; foot length 38.8% SVL;
relative lengths of toes I ,II ,III V,IV; heels without
tubercles; inner metatarsal tubercle weakly developed and
elongate; outer metatarsal tubercle absent; webbing formula
for the feet: I0–0II0–2III1–3IV2
–2V; no expanded pads
on toes.
FIG. 4.—Map of Sumatra with topographical relief showing localities where members of the new genus of toad (Sigalegalephrynus) were found.
70 Herpetologica 73(1), 2017
Color in life.—Edges of lore and head golden with black
shades; area below eyes with prominent white marking with
yellowish tint; dorsum light greenish-brown with light brown
hourglass figure extending from posterior of orbit to top of
the sacroiliac joint; hourglass shape ends with distinct
horizontal black bean color mark on each side; yellowish-
green marking on each shoulder; flanks black with red
tubercles, maculated with greenish-yellow blotches, and
possessing very prominent dark brown stripe starting from
posterior end of orbit to inguinal region; inguinal areas
greenish with golden tint; sacroiliac joint to inguinal region
of flanks, golden yellowish-green; dorsal sides of limbs light
brown; forearm, femur, tibia–fibula, and tarsus with distinct
dark spot encircled with golden-yellowish-green color;
venter opaque; throat golden-yellow; abdomen, ventral side
of arms and legs pinkish with scattered yellow and black
blotches; webbing in hand and foot translucent.
Color in preservative.—In preservative, the animal
appears dull because it has lost its golden-yellowish and
greenish colors. The hourglass pattern has turned gray. The
venter has lost all of its pinkish and golden-yellowish shades
and turned a greenish-white.
Etymology.—The specific epithet refers to the Minang-
kabau or Minang ethnic group inhabiting the region where
the new species was found.
Common name.—Minangkabau Puppet Toads.
FIG. 5.—Habitat of Sigalegalephrynus mandailinguensis—a view of the
rainforest at the edge of an inactive solphatara field on the northeastern
slope of Gunung Sorikmarapi where the holotype was found (upper); and
first author at the entrance of the subterranean hollow where the holotype
was collected (lower).
FIG. 6.—Advertisement call of Sigalegalephrynus mandailinguensis. (A)
oscillogram (relative amplitude vs. time) and (B) spectrograms (energy in
each frequency vs. time) of a single pulse; (C) power spectra. A color version
of this figure is available online.
Distribution and natural history.Sigalegalephrynus
minangkabauensis is known only from Gunung Kunyit from
an elevation of 1428 m (Fig. 4). The holotype was found
perched on a leaf ~1.25 m above ground, by the edge of a
forest stream at 2015 h. Before collecting it, ENS watched
the specimen move in reverse toward the edge of the leaf on
which it was perched, where it defecated (outside of the
surface of the leaf), and then return to its original position.
The holotype weighed 0.5 g.
Advertisement Calls
There is limited information on the acoustic properties of
advertisement calls for toads of Sundaland. Nevertheless, a
comparison with call data collected in previous field
expeditions and available information shows that the rapid
pulse of metallic ticks vocalized by Sigalegalephrynus
mandailinguensis is unique in its structure and spectral
properties among calls made by bufonids in the Sunda Shelf.
FIG. 7.—(A) Lateral, (B) dorsal, and (C) ventral view of the holotype of Sigalegalephrynus minangkabauensis from Gunung Kunyit, Sumatera Utara
Province, Sumatra (MZB 25738). A color version of this figure is available online.
72 Herpetologica 73(1), 2017
The advertisement call of Duttaphrynus melanostictus, for
example, is a shrill, gritty chatter that has been described as a
long train of pulses repeated at a regular rate, with an
average dominant frequency of 1293 kHz (Ngo and Ngo
2013). Ansonia leptopus calls have a varying range of
dominant frequency and, unlike S. mandailinguensis, consist
of two distinct pulses, one short and one long (Matsui 1982).
Phrynoidis asper advertisement calls are typically a three-
note repertoire consisting of a high-pitched ‘‘ bock’’ followed
by a couple of deep, raspy honks. The call of Leptophryne
borbonica usually consists of croaky crooning that is
reminiscent of marbles rubbing against each other. Ingero-
phrynus biporcatus calls are best described as a brisk series
of guttural bleats, whereas those of Pseudobufo subasper
consist of a volley of squeaky chirps. The pulses of the Indian
endemic Pedostibes tuberculosus have been described as
rapid series of schirr schirr, with a dominant frequency of
3782.13 kHz (Gururaja and Ramachandra 2006).
Phylogeny of Sunda Shelf Toad Genera
The systematics of Southeast Asian bufonids at the generic
level has been challenging, and the recent molecular
phylogenetic attempts at clarifying higher lever relationships
have generally failed to resolve relationships (Frost et al. 2006;
Matsui et al. 2007, 2010; Van Bocxlaer et al. 2009, 2010).
Consequently, the taxonomy of Southeast Asian toad genera
has been in flux (e.g., Chandramouli and Amarasinghe 2016).
Based on existing data, this lack of phylogenetic resolution
appears to stem primarily from the presence of short internal
braches. A phylogeny showing long branches interspersed
within a backbone of short internal branch lengths is usually
indicative of rapid radiations and is known to present a
significant challenge for phylogenetic inference (Whitfield and
Lockhart 2007; Whitfield and Kjer 2008; Rothfels et al. 2012).
According to several phylogenetic studies (e.g., Pramuk et al.
2008; Van Bocxlaer et al. 2010), the early evolutionary history
of bufonids involved an episode of rapid expansion and
radiation out of South America that allowed for a nearly
cosmopolitan distribution in a very short time. This could
explain the difficulty in achieving clearer resolution for the
deeper nodes in the phylogeny of South and Southeast Asian
toad genera. However, short internal nodes could also be
symptomatic of inadequate sampling, disagreement within or
among data sets, or loss of phylogenetic signal over time
(Whitfield and Lockhart 2007; Whitfield and Kjer 2008), in
which case, a more thorough sampling of taxonomic diversity
and selection of more informative loci would provide better
resolution for this clade. The relative influence of the
aforementioned factors should be addressed prior to making
any additional attempts at inferring the higher level
relationships of Sundaland bufonids.
Although patterns of the early diversification of Sundaland
bufonids remain poorly resolved in our study, the phylogeny
recovered by our analyses is, for the most part, congruent with
previous studies (Frost et al. 2006; Matsui et al. 2007; Van
Bocxlaer et al. 2009, 2010; Pyron and Wiens 2011). Ours is the
first study to incorporate the Bleeding Toads (Leptophryne
cruentata), whose generic affinity to L. borbonica, as
designated by morphology (Fitzinger 1843), is upheld by our
molecular results. Ours is also the first study to include the
unique aquatic toads (Pseudobufo subasper) in a phylogenetic
assessment. The relationship of a Duttaphrynus melanostictus
specimen from Sumatra with Pedostibes tuberculosus agrees
with Van Bocxlaer et al. (2009), who suggested Duttaphrynus
formed the sister taxon to Xanthophryne, an Indian endemic
not included in our study. This appears to indicate an Indian
origin for Duttaphrynus, with a recent invasion into Southeast
Asia. The lack of higher level phylogenetic resolution,
however, limits any attempt at a biogeographic appraisal of
these toads. In the context of the nonmonophyly of Pedostibes,
our results agree with Chan et al. (2016).
Based on the most comprehensive representation of
Sundaland toad genera to date, our results provide a
phylogenetic framework for future systematic and taxonomic
studies. More importantly, our analyses using both maximum
likelihood and Bayesian inference show the new taxa to be
phylogenetically isolated from all other genera with high
support, presenting a strong justification for the recognition
of a new genus.
Acknowledgments.—All specimens were collected and euthanized
following approved protocols (UTA IACUC A12.004). Research in Indonesia
was conducted under research permits149/SIP/FRP/SM/V/2013 (E.N. Smith)
and 155A/SIP/FRP/SM/XII/2013 (U. Smart). A National Science Foundation
grant (DEB-1146324) to ENS and MBH funded this research. We are
grateful to the Ministry of Research and Technology of the Republic of
Indonesia, RISTEK, for granting research permissions. S. Wahyono
(RISTEK) provided assistance throughout the permit approval process. We
are grateful to representatives of LIPI at the MZB for facilitating in-house
study of specimens and research permits, especially Boadi, R. Ubaidillah, and
Ir. R.M. Marwoto. RISTEK and LIPI approved our fieldwork in Indonesia
and provided export permits for specimen accessioning at UTA. W. Tri
laksono and A. Ryanto provided laboratory assistance at MZB. Mr. Widodo
and Marwoto from the Faculty of Mathematicas and Natural Sciences of
Universitas Brawijaya (MIPA-UB) provided logistical support. The Forestry
Department of Indonesia provided research permits for areas under their
jurisdiction: Kerinci Seblat NP (Sungai Penuh) and Batang Gadis NP
(Gunung Sorikmarapi). We thank Hartanto (DITJEN PHKA, Jakarta) for
help with forestry permits. We thank the local communities at Warkuk Ranau
Selatan (Sumatera Selatan) for their hospitality and logistical support. We
thank members of the Summer 2013 expedition to southern Sumatra: G.
Barraza (Broward College); W. Trilaksono (MZB); C. Franklin, K. O’Connell,
E. Wostl (UTA); and A.M. Kadafi, D.R. Wulandari, R. Darmawan, K.I.
Nawie, A. Dharasa, and S. Pratassi (MIP UB).
ABSTRAK (INDONESIAN): Kami mendeskripsikan satu
marga baru dan dua jenis baru kodok dari wilayah
gunung api di Provinsi Sumatera Utara (Gunung Sorik
Merapi) dan Provinsi Jambi (Gunung Kunyit). Takson
baru ini dapat dibedakan satu sama lain maupun
dengan marga kodok lainnya berdasarkan perbedaan
genetik, morfologi, dan struktur suara panggilan kodok
tersebut. Kami menggunakan data DNA mitokondria
dan DNA inti kodok-kodok dari famili Bufonidae yang
terdapat di wilayah Paparan Sunda untuk membuat
hipotesis filogenetik mengenai hubungan kekerabatan
antar marga kodok tersebut. Secara umum, hasil
penelitian yang kami peroleh mendukung hasil
penelitian-penelitian sebelumnya. Di samping itu,
kami juga menemukan untuk pertama kalinya bahwa
posisi filogenetik Pseudobufo dan marga baru berada di
bagian paling dasar dari seluruh marga katak di daerah
Paparan Sunda, dengan pengecualian pada marga
Alfaro, M.E., S. Zoller, and F. Lutzoni. 2003. Bayes or bootstrap? A
simulation study comparing the performance of Bayesian Markov chain
Monte Carlo sampling and bootstrapping in assessing phylogenetic
confidence. Molecular Biology and Evolution 20:255–266.
Audacity Team. 2014. Audacityt: Free audio editor and recorder, version
2.0.3. Available at Archived by WebCite
at on 6 December 2016.
Barbour, T. 1938. Notes on Nectophryne. Proceedings of the Biological
Society of Washington 51:191–196.
Bergsten, J. 2005. A review of long-branch attraction. Cladistics 21:163–193.
Biju, S., I. Van Bocxlaer, V.B. Giri, S.P. Loader, and F. Bossuyt. 2009. Two
new endemic genera and a new species of toad (Anura: Bufonidae) from
the Western Ghats of India. BMC Research Notes 2:241.
Castresana, J. 2000. Selection of conserved blocks from multiple alignments
for their use in phylogenetic analysis. Molecular Biology and Evolution
Chan, K.O., L.L. Grismer, A. Zachariah, R.M. Brown, and R.K. Abraham.
2016. Polyphyly of Asian tree toads, Genus Pedostibes unther, 1876
(Anura: Bufonidae), and the description of a new genus from Southeast
Asia. PLoS One 11:e0145903. DOI:
Chandramouli, S.R., and A.A.T. Amarasinghe. 2016. Taxonomic reassess-
ment of the arboreal toad Genus Pedostibes unther 1876 (Anura:
Bufonidae) and some allied Oriental bufonid genera. Herpetologica
Cocroft, R.B., and M.J. Ryan. 1995. Patterns of advertisement call evolution
in toads and chorus frogs. Animal Behaviour 49:283–303.
de Queiroz, K. 2005. A unified concept of species and its consequences for
the future of taxonomy. Proceedings of the California Academy of
Sciences 56:196–215.
Duellman, W.E. 2001. Hylid Frogs of Middle America. Society for the Study
of Amphibians and Reptiles, USA.
Duellman, W.E., A.B. Marion, and S.B. Hedges. 2016. Phylogenetics,
classification, and biogeography of the treefrogs (Amphibia: Anura:
Arboranae). Zootaxa 4104:1–109.
Fitzinger, L.J.F.J. 1843. Systema Reptilium. Fasciculus Primus. Braum¨uller
et Seidel, Austria. [In Latin.]
Frost, D.R., T. Grant, J. Faivovich, .. ., W.C. Wheeler. 2006. The amphibian
tree of life. Bulletinof the American Museum of Natural History 297:1–291.
Gururaja, K.V., and T.V. Ramachandra. 2006. Pedostibes tuberculosus
(Malabar Tree Toad) advertisement call and distribution. Herpetological
Review 37:75–76.
Hillis, D.M., and J.J. Bull. 1993. An empirical test of bootstrapping as a
method for assessing confidence in phylogenetic analysis. Systematic
Biology 42:182–192.
Huelsenbeck, J.P., and B. Rannala. 2004. Frequentist properties of Bayesian
posterior probabilities of phylogenetic trees under simple and complex
substitution models. Systematic Biology 53:904–913.
Kok, P.J., and M. Kalamandeen. 2008. Introduction to the Taxonomy of the
Amphibians of Kaieteur National Park, Guyana. Abc Taxa, Belgium.
Lanfear, R., B. Calcott, S.Y. Ho, and S. Guindon. 2012. PartitionFinder:
Combined selection of partitioning schemes and substitution models for
phylogenetic analyses. Molecular Biology and Evolution 29:1695–1701.
Larkin, M.A., G. Blackshields, N.P. Brown, R. Chenna, P.A. McGettigan, H.
McWilliam, F. Valentin, I.M. Wallace, A. Wilm, and R. Lopez. 2007.
Clustal W and Clustal X version 2.0. Bioinformatics 23:2947–2948.
Matsui, M. 1982. Call characteristics of several anuran species from East
Kalimantan. Contributions from the Biological Laboratory, Kyoto
University 26:131–139.
Matsui, M. 1984. (Genus Bufo, Bufonidae). Contributions from the
Biological Laboratory, Kyoto University 26:209–428.
Matsui, M., P. Yambun, and A. Sudin. 2007. Taxonomic relationships of
Ansonia anotis and Pedostibes maculatus, with a description of a new
genus (Amphibia, Bufonidae). Zoological Science 24:1159–1166.
Matsui, M., A. Tominaga, W. Liu, . . ., R.M. Brown. 2010. Phylogenetic
relationships of Ansonia from Southeast Asia inferred from mitochondrial
DNA sequences: Systematic and biogeographic implications (Anura:
Bufonidae). Molecular Phylogenetics and Evolution 54:561–570.
Miller, M.A., W. Pfeiffer, and T. Schwartz. 2010. Creating the CIPRES
Science Gateway for inference of large phylogenetic trees. New Orleans.
Pp. 1–8 in Proceedings of the Gateway Computing Environments
Workshop. New Orleans.
Mulcahy, D.G., T.H. Beckstead, and J.W. Sites, Jr. 2011. Molecular
systematics of the Leptodeirini (Colubroidea: Dipsadidae) revisited:
Species-tree analyses and multi-locus data. Copeia 2011:407–417.
Myers, C.W., and W.E. Duellman. 1982. A new species of Hyla from Cerro
Colorado, and other tree frog records and geographical notes from
western Panama. American Museum Novitates 52:1–25.
Ngo, B.V., and C.D. Ngo. 2013. Reproductive activity and advertisement
calls of the Asian common toad Duttaphrynus melanostictus (Amphibia,
Anura, Bufonidae) from Bach Ma National Park, Vietnam. Zoological
Studies 52:12.
Pramuk, J.B., T. Robertson, J.W. Sites, Jr., and B.P. Noonan. 2008. Around
the world in 10 million years: Biogeography of the nearly cosmopolitan
true toads (Anura: Bufonidae). Global Ecology and Biogeography 17:72–
Pyron, R.A., and J.J. Wiens. 2011. A large-scale phylogeny of Amphibia
including over 2800 species, and a revised classification of extant frogs,
salamanders, and caecilians. Molecular Phylogenetics and Evolution
R Core Team. 2013. R: A language and environment for statistical
computing, version 0.98.1062. R Foundation for Statistical Computing,
Austria. Available at Archived by WebCite at on 6 December 2017.
Rambaut, A. 2007. FigTree: A graphical viewer of phylogenetic trees.
Available at Archived by Web-
Cite at on 6 December 2017.
Rambaut, A., and A.J. Drummond. 2009. Tracer, version 1.5.0. Available at Archived by WebCite at http://www. on 6 December 2017.
Rohland, N., and D. Reich. 2012. Cost-effective, high-throughput DNA
sequencing libraries for multiplexed target capture. Genome Research
Ronquist, F., and J.P. Huelsenbeck. 2003. MrBayes 3: Bayesian phyloge-
netic inference under mixed models. Bioinformatics 19:1572–1574.
Rothfels, C.J., A. Larsson, L.Y. Kuo, P. Korall, W.L. Chiou, and K.M. Pryer.
2012. Overcoming deep roots, fast rates, and short internodes to resolve
the ancient rapid radiation of Eupolypod II Ferns. Systematic Biology
Sabaj Perez, M.H. (ed.). 2014. Standard Symbolic Codes for Institutional
Resource Collections in Herpetology and Ichthyology: An Online
Reference, Version 5.0 (22 September 2014). American Society of
Ichthyologists and Herpetologists, USA. Available at
Archived by WebCite at on 6
December 2017.
Stamatakis, A. 2006. RAxML-VI-HPC: Maximum likelihood-based phylo-
genetic analyses with thousands of taxa and mixed models. Bioinformatics
Stamatakis, A. 2014. RAxML Version 8: A tool for phylogenetic analysis and
post-analysis of large phylogenies. Bioinformatics 30:1312–1313.
Sueur, J., S. Pavoine, O. Hamerlynck, and S. Duvail. 2008. Rapid acoustic
survey for biodiversity appraisal. PLoS One 3:e0004065. DOI: http://dx.
Talavera, G., and J. Castresana. 2007. Improvement of phylogenies after
removing divergent and ambiguously aligned blocks from protein
sequence alignments. Systematic Biology 56:564–577.
Tamura, K., D. Peterson, N. Peterson, G. Stecher, M. Nei, and S. Kumar.
2011. MEGA5: Molecular evolutionary genetics analysis using maximum
likelihood, evolutionary distance, and maximum parsimony methods.
Molecular Biology and Evolution 28:2731–2739.
Van Bocxlaer, I., S.D. Biju, S.P. Loader, and F. Bossuyt. 2009. Toad
radiation reveals into-India dispersal as a source of endemism in the
Western Ghats-Sri Lanka biodiversity hotspot. BMC Evolutionary
Biology 9:131.
Van Bocxlaer, I., S.P. Loader, K. Roelants, S.D. Biju, M. Menegon, and F.
Bossuyt. 2010. Gradual adaptation toward a range-expansion phenotype
initiated the global radiation of toads. Science 327:679–682.
Whitfield, J.B., and K.M. Kjer. 2008. Ancient rapid radiations of insects:
Challenges for phylogenetic analysis. Annual Review of Entomology
Whitfield, J.B., and P.J. Lockhart. 2007. Deciphering ancient rapid
radiations. Trends in Ecology & Evolution 22:258–265.
Accepted on 8 November 2016
Associate Editor: Christopher Raxworthy
Specimens Examined
Ansonia glandulosa (n¼1).—INDONESIA: Sumatera Selatan: MZB
4239 (male).
74 Herpetologica 73(1), 2017
Ansonia latidisca (n¼1).—INDONESIA: Kalimantan Barat: Mount
Damus, Sambas, RMNH 10677–Holotype.
Ansonia spinulifer (n¼2).—INDONESIA: Kalimantan: Kabupaten
Singkawang: UTA 62129 (female), UTA 62130 (male).
Duttaphrynus melanostictus (n¼11).—INDONESIA: Jawa Barat:
Kabupaten Bandung: Gunung Kencana, 1751 m, 7.285968S, 107.638428E,
UTA 63437 (female); Cicayur, 839 m, 7.301998S, 107.577258E,UTA 63438
(male); road from Pangelengan to Sri Kandi, 1557 m, 7.258898S,
107.615548E, UTA 63439 (male); Kabupaten Bogor: Desa Sukamaki, P.T.
Vivaria indonesia, Frog Farm, 665 m, 6.6698338S, 106.9578338E, UTA
53738 (female); 6.6698338S, 106.9578338E, UTA 53739 (male); Taman
Safari, 1115 m, 6.7260008S, 106.9508338N, UTA 53740 (female) Hotel
Wisma Tamu, Kampus IPB, 192 m, 6.555568S, 106.726978E, UTA 62527
(male); Jawa Timur: Kabupaten Malang: Between Ngasem and Ngadilang-
kung, SE foothills of Gunung Butak, 359 m, 8.094818S, 112.567748E, UTA
62528 (male); road from Kraton Gunung Kawi to Malang, 360 m, 8.096448S,
112.53738E, UTA 62530 (female); Lampung: Kabupaten Lampung Selatan:
Gunung Rajabasa: S slope, 150–250 m, 5.8048338S, 105.6228338E, UTA
53746 (female); Kepulauan Riau: Bintan Island: RMNH 3961–Lectotype
Ingerophrynus biporcatus (n¼11).—INDONESIA: Jawa Barat:
Kabupaten Bogor: Kampus IPB pond, 225 m, 6.558778S, 106.720768E,
UTA 63768; Lampung: Kabupaten Lampung Selatan: Gunung RajabasPan-
pul Sukaraja Trail, 170 m, UTA 53731 (male); south slope Gunung Rajabasa,
Desa Canti, 70 m, 5.809378S, 105.597598E, UTA 63400 (female); 102 m,
5.807918S, 105.600788E, UTA A 63399 (female); 122 m, 5.801648S,
105.597728E, UTA 63401 (female); 31 m, 5.806138S, 105.595998E, UTA
63402 (female); Kabupaten Pasawaran: Margadalom, Padang Cermin, 10 m,
5.5541678S, 105.1858338E, UTA 53730; Hill’s North of Padang Cermin, 259
m, 5.523358S, 105.189968E, UTA 63403 (female); Gunung Pesawaran, 936
m, 5.517918S, 105.07318E, UTA 63406 (male); Kabupaten Tanggamus:
Vicinity of Ngarip Geothermal Plant, 777 m, 5.325368S, 104.577098E, UTA
63404 (male); Hill Above Ngarip, 1233 m, UTA 63405 (male).
Ingerophrynus claviger (n¼3).—INDONESIA: Sumatera Selatan:
Unknown: Road from Pagar Alam to Manna, 593 m, 4.185278S,
103.081598E, UTA 63770 (male), UTA 637701 (male), UTA 637702 (male).
Ingerophrynus parvus (n¼1).—INDONESIA: Sumatera Utara:
Kabupaten Deli Serdang: ‘‘Boy Scout Camp’’ in Bandar Baru, 904 m,
3.268358N, 98.539538E, UTA 63769 (male).
Leptophryne borbonica (n¼16).—INDONESIA: Bengkulu: Unknown:
MZB 2931; MZB 3479 (female); MZB 3554 (male); MZB 9337 (male); MZB
16312 (female); Jambi: Unknown: MZB 21845 (male); Jawa: Unknown:
MZB 23838 (male); MZB 6716 (female); MZB 6717 (male); MZB 6719
(female); RMNH 1739–Syntype; Jawa Barat: MZB 15155 (female);
Kalimantan: Unknown: MZB 3641 (male); Lampung: Kabupaten Tangga-
mus: MZB 3445 (female); hill above Ngarip, 5.282188S, 104.557738E, UTA
62486; Sumatera Selatan: Unknown: MZB 4234 (female).
Leptophryne cruentata (n¼2).—INDONESIA: Jawa Barat: Kabupaten
Cianjur: Gunung Gede Pangrango National Park, 1421 m, 6.743458S,
107.003558E, UTA 62522; Cibodas, RMNH 2130–Syntype.
Pelophryne brevipes (n¼6).—INDONESIA: Jambi: MZB 14950
(female); Lampung: Kabupaten Pesawaran: Gunung Pesawaran, 1046 m,
5.515638S, 105.076678E, UTA 63762; Unknown: MZB 14607; Sumatera
Barat: MZB 3185 (male); Sumatera Selatan: MZB 12078 (female);
Unknown: Unknown: MZB 17097 (female).
Pelophryne signata (n¼6).—INDONESIA: Jambi: MZB 21.864
(female); MZB 21.863; Kalimantan Barat: MZB 4342 (male); MZB 4343
(male); MZB 4344 (female); Kalimantan Timur: MZB 6283 (female).
Phrynoides asper (n¼10).—INDONESIA: Jawa Barat: Kabupaten
Bogor: Desa Sukamaki P.T. Vivaria indonesia, Frog Farm, 660 m,
6.6698338S, 106.9578338E, UTA 53719; Lampung: Kabupaten Lampung
Barat: Kubuperahu, 730 m, 5.064678S, 104.036238E, UTA 62469 (female);
Kabupaten Pasawaran: Road between Padang Cermin and Kedongdong, 382
m, 5.546538S, 105.046788E, UTA 61449 (male); road between Padang
Cermin and Kedongdong 454 m, 5.551968S 105.061318E, UTA 62448
(male); road between Padang Cermin and Kedongdong 189 m, 5.487878S,
105.016858E, UTA 62453 (male), UTA 62454 (male); Kabupaten Tangga-
mus: Ngarip paddy fields, 869 m, 5.306388S, 104.548278E, UTA 62459
(male); forest above Ngarip Town, 1324 m, 5.28228S, 104.556938E, UTA
62463 (female); NE of town of Ngarip, 830 m, 5.314468S, 104.540948E, UTA
62467; Unknown: Unknown: RMNH 2172–Lectotype.
Phrynoides juxtasper (n¼9).—INDONESIA: Jambi: Kabupaten
Kerinci: beginning of trail to Danau Tujuh, 1409 m, 2.041398S,
101.314628E, UTA 62414 (female); trail to Danau Tujuh, 1565 m,
1.708688S, 101.369818E, UTA 62415 (male); Unknown: Unknown: Road
between Tapan and Sungaipenuh, 30 km W of Sungaipenuh, UTA 63767
(male); Jawa Barat: Kabupaten Bogor: Desa Sukamaki P.T. Vivaria
Indonesia, Frog Farm, 660 m, 6.6720008S, 106.8796678E, UTA 53719
(female); Sumatera Utara: Kabupaten Deli Serdang: Boy Scout Camp in
Bandar Baru, 901 m, 3.268988N, 98.539928E, UTA 63431 (male); Boy Scout
Camp in Bandar Baru, 906 m, 3.267878N, 98.539678E, UTA 63432 (female);
Kabupaten Samosir: W coast of Island S of Pangururan, 926 m, 2.456818N,
98.84388E, UTA 63434 (male); Kabupaten Mandailing Natal: Kota Baringen
Julu (Batang Gadis National Park), 1174 m, 0.666088N, 99.578938E, UTA
63436 (female); Kalimantan Timur: Unknown: Upper Mahakam River,
RMNH 14517–Paratype.
Pseudobufo subasper (n¼4).—INDONESIA: Riau: Sungai Kerumutan,
UTA 63763 (female); UTA 63764 (male); Unknown: Unknown: RMNH
2199–Syntype (male); RMNH 2200–Syntype (female); RMNH 2216–
Rentapia hosii (n¼9).—UNKNOWN: Unknown: Unknown: UTA 38040
(female); UTA 43516 (female); UTA 6395 (female); UTA 6396 (female);
UTA 6430 (female); UTA 6436 (female); UTA 6494 (male); UTA 6520
(male); UTA 6705 (female).
Rentapia rugosa (n¼1).—MALAYSIA: Sarawak: MZB 8401.
Sigalegalephrynus mandailinguensis (n¼4).—INDONESIA: Sumatera
Utara: Kabupaten Mandailing Natal: Gunung Sorikmarapi (W side), 1383 m,
0.701648N, 99.552628E, MZB 25736–Holotype (male); trail between the
Tano Bato to Sapo Tinjak road and Lake Saba Begu, Batang Gadis National
Park, 1297 m 0.708668N, 99.519538E, MZB 25737–Paratype (male); 1299 m,
0.708668N, 99.519538E, UTA 63562–Paratype (male), UTA 63561–Paratype
Sigalegalephrynus minangkabauensis (n¼1).—INDONESIA: Jambi:
Kabupaten Kerinci: Gunung Kunyit, 1428 m, 2.260138S, 101.495128E, MZB
... Even though 24% of the amphibian species have been categorized as Data Deficient (DD) (Nori et al. 2018), about 80% do not have their up-to-date IUCN Red List status (Tapley et al. 2018) (many in Malaysia, Indonesia, Papua New Guinea, China). To help prioritize conservation for the newly discovered arboreal Puppet Toads, genus Sigalegalephrynus Smart, Sarker, Arifin, Harvey, Sidik, Hamidy, Kurniawan & Smith (2017), we describe three newly discovered populations as new species and provide IUCN Red List status assessment for all members of this Sumatran endemic genus. ...
... The genus Sigalgalephrynus contains two species: S. mandailinguensis, from Gunung Sorikmerapi, Batang Gadis National Park, Sumatera Utara province, and S. minangkabauensis, from Gunung Kunyit, a peak in the Barisan Range of the province of Jambi (Smart et al. 2017). In 2015, we collected additional specimens of this genus, from the Burni Telon volcano and Highlands of Gayo Lues in the province of Aceh, and from Dempo volcano of the Sumatera Selatan province. ...
... For a very narrow-ranged species, this minimum number of occurrence data points can be as low as three (Proosdij et al. 2016) since MaxEnt is less sensitive to sample size (Wisz et al. 2008). Sigalegalephrynus species are micro-endemics (Smart et al. 2017) and occurrence data of these is limited, in fact we have no more than two useful occurrence data points for a single species. Thus, for finding suitable habitats for potential new populations or species, instead of modeling each species individually we collectively modeled our nine GPS data points for the five Sigalegalephrynus species. ...
Full-text available
Using a combination of morphological and molecular data we recognize three new species of Puppet Toad, Sigalegalephrynus Smart, Sarker, Arifin, Harvey, Sidik, Hamidy, Kurniawan & Smith, a recently described genus endemic to the highland forests of Sumatra, Indonesia. Phylogenetic analysis of mitochondrial DNA sequences recovered a monophyletic relationship among all Puppet Toads, with two distinct evolutionary clades, a northern and a southern. The northern clade includes Sigalegalephrynus gayoluesensis sp. nov., and S. burnitelongensis sp. nov., and the southern clade includes S. harveyi sp. nov., S. mandailinguensis, and S. minangkabauensis. With the discovery of these three new species, Sigalegalephrynus contains more endemic species than any other genus of toad in Indonesia. We used maximum entropy, implemented in MaxEnt, to identify suitable habitats and occurrence probability of additional undescribed new species from the island. The most important predictors of Sigalegalephrynus distribution were elevation (64.5%) and land cover (7.11%). Based on the probability of presence, it is likely that there are many more species of the genus awaiting discovery in Sumatra. Our analysis, based on IUCN Red List of Threatened Species category and criteria, shows that all of the five species of Sigalegalephrynus are in great risk of extinction and should be placed into the Endangered (EN) category of IUCN Red List.
... Although our molecular analysis was based upon a relatively short fragment of a single gene, our MLBS and NJBS analyses supported the genus Leptophryne to be monophyletic group, congruent the findings of Smart et al. (2017) who used both mitochondrial and nuclear genes. By examining type specimens and newly collected topotypic material we were able to elucidate for the first time the phylogenetic position of L. borbonica, L. cruentata and L. javanica sp. ...
... Our results also revealed that populations of L. borbonica from Borneo and Sumatra (Sumatra I and II) form distinct molecular lineages, separated from true L. borbonica (west Java/ topotype). The two lineages from Sumatra (Sumatra I from Bengkulu and Sumatra II from Lampung) are separated by 300 km, and by the Bukit Barisan Selatan mountain ranges, which may form a barrier to dispersal, as has been shown in frog species in the genus Rhacophorus (Harvey et al. 1996;Streicher et al. 2014;Hamidy & Kurniati 2015), Philautus (Wotzl et al. 2017), Chiromantis (Riyanto & Kurniati 2012), Sigalegalephrynus (Smart et al. 2017), Sumaterana (Arifin et al. 2018) and the Limnonectes kuhlii group (McLeod et al. 2011). It is likely that further undiagnosed diversity exists within the genus, and more comprehensive examination of acoustic data and detailed morphological data are likely to result in Sumatran and the Bornean populations being recognised as distinct lineages. ...
We investigated phylogenetic relationships among populations of two species within the genus Leptophryne, L. cruentata and L. borbonica, using the mitochondrial 16S rRNA gene. As a result, we identified two distinct lineages within populations currently considered Leptophryne cruentata: 1) a lineage containing L. cruentata from the type locality, and 2) a lineage from Mt. Slamet and Mt. Ciremai. On the basis of genetic and morphological differences, we describe the Mt. Slamet and Mt. Ciremai populations as a new species, L. javanica sp. nov. The new species is distinguished from L. cruen-tata and L. borbonica by the following combination of morphological characters: the presence of distinct yellow mottling on the dorsum; relatively small body size (SVL male 22.2-24.0 mm, female 29.6 mm); relatively short hindlimbs (HLL 37.0-40.9 mm); relatively short fourth toe (4ToeL 4.3-6.0 mm); basal webbing on the hands, but well developed on toes; very protruding snout and an indistinct tympanum. In our preliminary phylogenetic analysis, we also detected four distinct lineages within Leptophryne borbonica: 1) a lineage containing true L. borbonica from west Java, 2) a lineage from Lam-pung (Sumatra I), 3) a lineage from northern Borneo, and 4) a lineage from Bengkulu (Sumatra II). Further studies are needed to determine taxonomic status of these lineages.
... Herpetofauna discoveries began primarily in Java in the nineteenth century (Boulenger 1890, de Rooij 1917, van Kampen 1923, Kopstein 1930, de Haas 1941 and expedited our understanding of Indonesia's species diversity. Recent decades have seen a rapid increase in the number of newly described herpetofauna species, particularly for amphibians, e.g., Megophrydae (Hamidy and Matsui 2010, Eto et al. 2018, Rhacophoridae , Hamidy and Kurniati 2015, Wostl et al. 2017a, Mediyansyah et al. 2019, Microhylidae (Matsui et al. 2013, Atmaja et al. 2019, Munir et al. 2020, Eprilurahman et al. 2021, Bufonidae (Smart et al. 2017, Sarker et al. 2019, Ranidae Hamidy 2012, Arifin et al. 2018), Dicroglossidae (Iskandar et al. 2011a, Mcleod et al. 2011; and reptiles, e.g., Gekkonidae including Cyrtodactylus (Iskandar 2011b, Riyanto et al. 2018a, Riyanto et al. 2018b, Cnemaspis (Amarasinghe et al. 2015a, Riyanto et al. 2017, Hemiphyllodactylus (Grismer et al. 2014), and Lepidodactylus (Stubbs et al. 2017); Agamidae (Harvey et al. 2014, Harvey et al. 2017a, Harvey et al. 2018, Harvey et al. 2019, Colubridae (Vogel et al. 2014, Amarasinghe et al. 2015b, Wostl et al. 2017a, Amarasinghe et al. 2021, and Cylindrophiidae (Kiecksbusch et al. 2016); which highlight the fact that many enigmatic, elusive, and cryptic species are waiting to be discovered. Although the region of Sumatra, Borneo, Celebes, Moluccas, and Java has been extensively surveyed (David and Vogel 1996, Inger et al. 2005, de Lang and Vogel 2006Kusrini et al. 2021), the primary focus has been on areas with relatively high diversity. ...
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An inventory of herpetofauna species from western part of Nusa Kambangan Island, Central Java, Indonesia, is presented. There are 43 herpetofauna species reported (16 amphibians and 27 reptiles). This study confirmed new distribution record and list some of threatened species. In light of the imminent human disturbances on Nusa Kambangan Island, a conservation plan is urgently needed.
... Between 2016 and 2020, 780 new species of amphibians were described (figure 3), a higher number than in the previous five years (732 species between 2011 and 2015). Most (705) of the new species described in the last five years were from the largest amphibian order, Anura (salamanders gained 66 species and caecilians gained 9). Six genera of frogs were described based entirely on newly discovered species: Astrobatrachus (Vijayakumar et al. 2019) (also representing the new subfamily Astrobatrachinae); Blythophryne (Chandramouli et al. 2016); Mini (Scherz et al. 2019) (figure 3f); Siamophryne ; Sigalegalephrynus (Smart et al. 2017); and Vietnamophryne . At a regional level, most new species added between 2016 and 2020 are from Latin America (40.8%), ...
Full-text available
Amphibians are a clade of over 8,400 species that provide unique research opportunities and challenges. With amphibians undergoing severe global declines, taking stock of our current understanding of amphibians is imperative. Focusing on 2016–2020, we assessed trends in amphibian publishing, conservation research, systematics, and community resources. We show that while research and data availability are increasing rapidly, information is not evenly distributed across research fields, clades, or geographic regions, leading to substantial knowledge gaps. A complete review of amphibian NCBI resources indicates that genomic data are poised for rapid expansion, but amphibian genomes pose significant challenges. A review of recent conservation literature and cataloged threats on 1,261 species highlight the need to address land use change and disease using adaptive management strategies. We underscore the importance of database integration for advancing amphibian research and conservation and suggest other understudied or imperiled clades would benefit from similar assessments.
... Most of these are considered to be allopatric with their closest relatives (Grismer et al., 2019;Matsui et al., 2017;Poyarkov et al., 2019;Chan et al., 2020a,b) but some have been found to be sympatric within the range of their close relatives, even if restricted to one or two mountains (Grismer et al., 2015;Eto et al., 2016;Quah et al., 2020). The highlands of Southeast Asia have high species diversity and endemism of amphibians and reptiles (Malkmus et al., 2002;Grismer et al., 2012;Karin et al., 2016;Smart et al., 2017;Arifin et al., 2018;Shaney et al., 2020). However, their diversity is threatened by habitat destruction and climate change (Bickford et al., 2010;Margono et al., 2014;Harris et al., 2017). ...
The genus Kalophrynus is represented by tiny to medium terrestrial or subfossorial frogs and is widely distributed in Southeast Asia. The diversity centre of this genus is Borneo, where almost half of all nominal species are distributed and all of these are endemic to this and the peripheral islands. Kalophrynus nubicola is the smallest species within this genus; it is found only at high elevations in Gunung Mulu National Park, northeastern Sarawak, Malaysian Borneo. Three groups of populations were preliminary reported, but taxonomic decision on the groups was not yet given. In this study, we investigated the morphological, acoustic, and genetic differences among these three populations to reassess their taxonomic status. Morphological analyses confirmed differences in body size and colouration, molecular analyses indicated that each group was monophyletic, and differences in the acoustic characteristics of each group were also diagnostic. Therefore, herein, we describe them as distinct taxa. Phylogenetic analyses revealed that the K. nubicola group forms a highly divergent clade from other species within the genus. Our findings reveal that seven Kalophrynus species occur in Gunung Mulu National Park, with non-overlapping distributions according to elevation or forest type. We further discuss the biogeography and evolutionary history of this species group.
... Pekerjaan tersebut menghasilkan beberapa deskripsi spesies baru, genus baru, deskripsi ulang spesies dan penemuan spesies yang hilang selama bertahun-tahun. Sebagai contoh deskripsi spesies baru (Wostl et al, 2017;Teynie et al, 2010;Harvey et al, 2017a,b;Munir et al, 2018;Atmaja et al, 2019); genus baru dengan beberapa spesies baru (Arifin et al, 2018;Smart et al, 2017); deskripsi ulang spesies (Harvey et ...
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The objective of this study is to invent amphibians inhabiting the region of natural tourism object of Malibo Anai. The region is located in the foot of mountain Tandikek in Sumatera Barat province. Since this resort serve as natural tourism object, the resort maintains primary forest existed in most of the area which might provide an appropriate home for amphibians and other group of animals. The survei was conducted in January and July 2020. We recorded 22 species of amphibians in three types of habitat. Most of amphibians were found near the standing body water. A species of Rhacophoridae, Zhangixalus achantharrhena and of Ranidae, Pulchrana fantastica were found and reported formally for the first time outside of their type localities. Along with this study, we also recorded snakes and several cryptic bent-toed geckoes. The area of study would serve as a good place to conduct research on biology and ecology.
... Similar to most bufonids in Southeast Asia and Southern China (Grosjean & Dubois 2001;Sukumaran et al. 2010;Matsui et al. 2017;Amram et al. 2018;Matsui 2019), breeding males of P. scalpta only produce a single type of advertisement call. The advertisement calls of P. scalpta is different from most known calls of bufonids in the region by having a moderate pitch (3.0-3.2 kHz): the advertisement calls are distinctly lower in large-sized toads of the genus Bufo (0.4 kHz, Chen et al. 2018), Duttaphrynus Frost et al. (1.3-1.6 kHz, Grosjean & Dubois 2001), Rentapia Chan, Grismer, Zachariah, Brown & Abraham (1-1.6 kHz, Amram et al. 2018;Chan et al. 2020) and Phrynoidis Fitzinger (0.9-2 kHz, Grosjean & Dubois 2001;Amram et al. 2018), distinctly higher in the genus Pelophryne (5.3-6.8 kHz, Matsui et al. 2017;Matsui 2019), moderate in the genus sigalegalephrynus Tschudi (3.4 kHz, Smart et al. 2017), and show great interspecific variations in the genus Ansonia Stoliczka (2.5-8.2 kHz, Sukumaran et al. 2010;Amram et al. 2018;Ong & Das 2019) (See Table 2). The causes of variations in dominant frequency among these bufonoid genus warrants further investigation. ...
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The Hainan Pygmy Toad Parapelophryne scalpta (Liu & Hu) is the single species of the genus Parapelophryne Fei, Ye & Jiang in the Family Bufonidae (Frost 2020) and is endemic to China’s Hainan Island, which lies within the tropics at 18°09′–20°10′N, 108°37′–111°03′E, with a land area of ca. 33,900 km2. The taxonomy status of this species has obtained much debates since its discovery: it was originally described by Liu et al. (1973) as a member of the genus Nectophryne Buchholz & Peters then allocated to Pelophryne Barbour by Ye & Fei (1978). On the basis of its unique morphological characters, Fei et al. (2003) erected a new genus Parapelophryne to discriminate P. scalpta from other bufonids. Some researchers, however, still listed the species under the genus Pelophryne (Stuart et al. 2008; Shi et al. 2011). Recently, the validity of the genus Parapelophryne is further confirmed by Matsui et al. (2015) on the basis of molecular analyses. In- terestingly enough, Matsui et al. (2015)’s result showed that this extremely small-sized toad is a sister taxon of the large-sized toads of the genus Bufo Garsault, which is mainly distributed in temperate and subtropical regions of Eurasia (Frost 2020). Parapelophryne scalpta is a forest-dependent species and only inhabits in well-preserved evergreen broadleaf forests (Fei & Ye 2016). It is the smallest toad species in China with snout-vent length of 19–23 mm in adult males and 24–27 mm in females (Fei & Ye 2016). Due to their small size and elusive habits, little is known about the natural history of P. scalpta and its acoustic characteristics are still undocumented (Fei & Ye 2016). During a herpetological survey in Hainan in 2015, we luckily detected male calls of P. scalpta and obtained a short but clear record of the advertisement call of the species. Herein, we describe the acoustic characteristics and calling behavior of Parapelophryne scalpta.
... Some researchers indicated that Sumateran amphibian and reptile inventories are limited [1,2,3,4]. However, some species were described as new taxa [4,5,6,7,8,9,10,11,12]. A survey has been conducted in August 2019 in mount Sago from the elevation of 1000 meter above sea level (masl) to 2567 masl. ...
... Singapore: Bukit Timah Nature Reserve (Lim and Lim, 1992). Indonesia: Lampung, Sumatra (Smart et al., 2017), ? Natuna Islands, ?Mentawei Islands (Inger, 1966). ...
A small, semi-arboreal toad of the genus Pelophryne from Peninsular Malaysia has been treated as P. brevipes or P. signata. The peninsular toad and Bornean P. signata are very similar to each other morphologically, although slightly different in relative forelimb length, dorsal coloration, and tuberculation. However, in partial mtDNA sequence, the peninsular toad is substantially distinct from P. signata from Borneo and P. brevipes from the Philippines, although it is close to a congener from Sumatra. Thus, the peninsular toad is described as a new species based on specimens from Genting Highlands, state of Pahang, central Peninsular Malaysia. Of the two morphotypes recognized in the genus, the new species belongs to the one with the tips of the fingers expanded into truncate discs, in which the new species is the smallest in body size. The new species also occurs in Singapore and possibly in Sumatra.
Aim The diversity of brood size across animal species exceeds the diversity of most other life‐history traits. In some environments, reproductive success increases with brood size, whereas in others it increases with smaller broods. The dominant hypothesis explaining such diversity predicts that selection on brood size varies along climatic gradients, creating latitudinal fecundity patterns. Another hypothesis predicts that diversity in fecundity arises among species adapted to different microhabitats within assemblages. A more recent hypothesis concerned with the consequences of these evolutionary processes in the era of anthropogenic environmental change predicts that low‐fecundity species might fail to recover from demographic collapses caused by rapid environmental alterations, making them more susceptible to extinctions. These hypotheses have been addressed predominantly in endotherms and only rarely in other taxa. Here, we address all three hypotheses in amphibians globally. Location Global. Time period Present. Major taxa studied Class Amphibia. Methods Using a dataset spanning 2,045 species from all three amphibian orders, we adopt multiple phylogenetic approaches to investigate the association between brood size and climatic, ecological and phenotypic predictors, and according to species conservation status. Results Brood size increases with latitude. This tendency is much stronger in frogs, where temperature seasonality is the dominant driver, whereas salamander fecundity increases towards regions with more constant rainfall. These relationships vary across continents but confirm seasonality as the key driver of fecundity. Ecologically, nesting sites predict brood size in frogs, but not in salamanders. Finally, we show that extinction risk increases consistently with decreasing fecundity across amphibians, whereas body size is a “by‐product” correlate of extinction, given its relationship with fecundity. Main conclusions Climatic seasonality and microhabitats are primary drivers of fecundity evolution. Our finding that low fecundity increases extinction risk reinforces the need to refocus extinction hypotheses based on a suggested role for body size.
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We reassessed the taxonomic status of an Asian genus of arboreal bufonids, Pedostibes, based on examination of preserved material of the two species currently attributed to this genus. Analysis of their morphological, morphometric, and geographic distribution data revealed that Pedostibes tuberculosus, the type species of this genus from the Western Ghats, southwestern India, is morphologically distinct from the geographically separated member, P. kempi, which is distributed in northeastern India. Hence, the generic nomen Pedostibes is restricted to the type species, rendering it a monotypic genus from the Western Ghats of peninsular India. A re-examination and detailed comparisons of the types of P. kempi with other bufonid genera revealed morphological similarities with another geographically proximate toad, Bufoides meghalayanus, from northeastern India. Hence, this taxon is formally transferred herein to Bufoides with a redescription. The composition of the recently described Southeast Asian toad genus Rentapia is reevaluated and the name-bearing type specimens of the currently ascribed taxa are redescribed. A detailed examination of the types of Rentapia everetti and R. rugosa revealed morphological congruence coupled with geographic sympatry. Hence, the latter nomen is synonymized with R. everetti in accordance with the International Code of Zoological Nomenclature principle of priority.
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The Asian Tree Toad genus Pedostibes, as currently understood, exhibits a conspicuouslydisjunct distribution, posing several immediate questions relating to the biogeography andtaxonomy of this poorly known group. The type species, P. tuberculosus and P. kempi, areknown only from India, whereas P. hosii, P. rugosus, and P. everetti are restricted to SoutheastAsia. Several studies have shown that these allopatric groups are polyphyletic, withthe Indian Pedostibes embedded within a primarily South Asian clade of toads, containingthe genera Adenomus, Xanthophryne, and Duttaphrynus. Southeast Asian Pedostibes onthe other hand, are nested within a Southeast Asian clade, which is the sister lineage to theSoutheast Asian river toad genus Phrynoidis.We demonstrate that Indian and SoutheastAsian Pedostibes are not only allopatric and polyphyletic, but also exhibit significant differencesin morphology and reproductive mode, indicating that the Southeast Asian species’are not congeneric with the true Pedostibes of India. As a taxonomic solution, we describe anew genus, Rentapia gen. nov. to accommodate the Southeast Asian species.
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We examined differences in reproductive activities and intraspecific variations in advertisement calls of Duttaphrynus melanostictus (Schneider, 1799) that lives in a tropical region of central Vietnam. The snout-vent length (SVL) of sexually mature males ranged from 50.2 to 70.3 mm, while that of females ranged from 65.2 to 97.3 mm. Histological analyses of testes revealed that sperm was not present throughout the year, indicating discontinuous reproductive activity for adult males. Adult females were captured year-round, but reproductive females were observed only during months of the auxiliary rainy season (mainly April to July), indicating that females reproduce seasonally. We also estimated levels of within-male variation of each call property and the influences of the ambient temperature, humidity, and the SVL of calling males on acoustic features. The call rate and pulse rate showed intermediate levels of variation, whereas the dominant frequency and call duration were the most stereotyped properties. One-way analyses of variance for six acoustic properties showed that the call rate, pulse rate, and rise time significantly differed (p < 0.05) among localities. Advertisement calls were a series of groups of 56 to 244 (145 ± 54) pulses with an average call duration of 26.722 s, an average pulse rate of 11.69 pulses/s, and an average dominant frequency of 1.293 kHz. The results of the multiple regressions for possible effects of temperature, humidity, and SVL on the six acoustic properties indicated that the dominant frequency, pulse rate, call duration, and rise time were positively significant. In Bach Ma National Park, when air temperature in the recording area decreased to <16.6°C, and advertisement calls of adult males virtually ceased in all three populations.
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Cat-eyed snakes (Leptodeira) were thought to be closely related to nightsnakes (Hypsiglena and Pseudoleptodeira) based on morphology and immunological data, which allied these genera with blunt-headed vine snakes (Imantodes) and the Cloud Forest Snake (Cryophis hallbergi). We collected sequence data from six protein-encoding nuclear loci (SLC30A1, ZEB2, FSHR, NTF3, DNAH3, and PNN; 4149 bp) and additional mtDNA data (nad5; 955 bp) added to published cob and nad4 (total 2387 bp mtDNA) from these and other rear-fanged, mildly venomous snakes that prey on vertebrates (frogs and lizards) and from several other dipsadine genera (Dipsas, Sibon, and Atractus) that prey on invertebrates (goo-eaters). We analyzed relationships using concatenation and a coalescent species-tree method. When analyzed separately, using either concatenation or coalescent methods, nuclear data support a different overall topology from the mtDNA data. Like the mtDNA data, the nuclear data support the Leptodeira + Imantodes relationship, but instead place this clade more closely to the goo-eaters, with the nightsnakes as the basal divergence in the group. When the data are combined in concatenation analyses, the more variable mtDNA data appear to overwhelm the nuclear data, but not under the coalescent model.
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Phylogenies are increasingly used in all fields of medical and biological research. Moreover, because of the next generation sequencing revolution, datasets used for conducting phylogenetic analyses grow at an unprecedented pace. RAxML (Randomized Axelerated Maximum Likelihood) is a popular program for phylogenetic analyses of large datasets under maximum likelihood. Since the last RAxML paper in 2006, it has been continuously maintained and extended to accommodate the increasingly growing input datasets and to serve the needs of the user community. I present some of the most notable new features and extensions of RAxML, such as, a substantial extension of substitution models and supported data types, the introduction of SSE3, AVX, and AVX2 vector intrinsics, techniques for reducing the memory requirements of the code and a plethora of operations for conducting post-analyses on sets of trees. In addition, an up-to-date, 50 page user manual covering all new RAxML options is available. The code is available under GNU GPL at
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Kaieteur National Park is a protected area covering ca. 63,000 ha located at the eastern edge of the Pakaraima Mountains, in a largely unexplored region of west-central Guyana. Next to providing description of the area, its vegetation and climate, an overview of the equipment and appropriate techniques needed to study amphibian taxonomy, this manual also provides a brief summary of our current knowledge of the amphibian systematics in the region, key features useful to identify amphibians, and the very first field guide dealing with the amphibian fauna of Guyana, notably with the amphibians of Kaieteur National Park. A total of 48 species (46 anurans and 2 caecilians) are treated and illustrated in colour. Field keys, field identifications, brief information on natural history, calls, tadpoles and distribution within and outside the Park are also included. This work also reports the microhylid Synapturanus salseri Pyburn, 1975 for the first time from Guyana
A phylogenetic analysis of sequences from 503 species of hylid frogs and four outgroup taxa resulted in 16,128 aligned sites of 19 genes. The molecular data were subjected to a maximum likelihood analysis that resulted in a new phylogenetic tree of treefrogs. A conservative new classification based on the tree has (1) three families composing an unranked taxon, Arboranae, (2) nine subfamilies (five resurrected, one new), and (3) six resurrected generic names and five new generic names. Using the results of a maximum likelihood timetree, times of divergence were determined. For the most part these times of divergence correlated well with historical geologic events. The arboranan frogs originated in South America in the Late Mesozoic or Early Cenozoic. The family Pelodryadidae diverged from its South American relative, Phyllomedusidae, in the Eocene and invaded Australia via Antarctica. There were two dispersals from South America to North America in the Paleogene. One lineage was the ancestral stock of Acris and its relatives, whereas the other lineage, subfamily Hylinae, differentiated into a myriad of genera in Middle America.