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The generic assignment of the draconine lizard Gonocephalus robinsonii from the highlands of West-Malaysia has been uncertain since the original description. Here we present a study based on morphology, previously published karyotype data and molecular phylogenetics using 16S rRNA sequences to evaluate the systematic status of G. robinsonii. As a result we describe Malayodracon gen. nov. to accommodate the species.
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Accepted by S. Carranza: 28 Sept. 2015; published: 3 Nov. 2015
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The systematic status of Gonocephalus robinsonii Boulenger, 1908
(Squamata: Agamidae: Draconinae)
WOLFGANG DENZER
1,5
, ULRICH MANTHEY
2
, KRISTIN MAHLOW
3
& WOLFGANG BÖHME
4
1
Society for Southeast Asian Herpetology, Calle Rio Segura 26, 30600 Archena, Murcia, Spain. E-mail: wolfdenoxford@yahoo.co.uk
2
Society for Southeast Asian Herpetology, Kindelbergweg 15, 12249 Berlin, Germany. E-mail: manthey.sseah@t-online.de
3
Museum für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, Ger-
many. E-mail: kristin.mahlow@mfn-berlin.de
4
Zoologisches Forschungsmuseum Alexander Koenig, Leibniz-Institut für Biodiversität der Tiere, Adenauerallee 160, 53113 Bonn,
Germany. E-mail: w.boehme@zfmk.de
5
Corresponding author. E-mail: wolfdenoxford@yahoo.co.uk
Abstract
The generic assignment of the draconine lizard Gonocephalus robinsonii from the highlands of West-Malaysia has been
uncertain since the original description. Here we present a study based on morphology, previously published karyotype
data and molecular phylogenetics using 16S rRNA sequences to evaluate the systematic status of G. robinsonii. As a result
we describe Malayodracon gen. nov. to accommodate the species.
Key words: Gonocephalus, phylogeny, cranial morphology, Malayodracon gen. nov., Malayodracon robinsonii comb.
nov., Dendragama boulengeri
Introduction
The genus Gonocephalus Kaup, 1825 constitutes a group of arboreal agamid lizards belonging to the subfamily
Draconinae distributed in the Sunda archipelago west of the Wallace line, the Philippines and mainland Southeast
Asia south of the Isthmus of Kra (with the exception of isolated populations of G. grandis (Gray, 1845) in southern
Laos [Teynet al. 2004] and Vietnam [Ananjeva et al. 2007]).
For a long time the genus Gonocephalus was a conglomerate of arboreal agamid lizards that comprised species
from Southeast Asia, Andaman & Nicobar Islands, the Sunda Archipelago, the Philippines, New Guinea, the
Bismarck Archipelago and Australia. Currently only those species from west of the Wallace line are considered
true Gonocephalus. The isolated population from Andaman and Nicobar Islands belong to the genus
Coryphophylax Blyth, 1860 and those species east of the Wallace line are members of the genus Hypsilurus Peters,
1867. These taxonomic changes had already been proposed by Moody (1980) based on phenotypic characters and
results of his phylogenetic study were corroborated through discovering additional characters separating
Gonocephalus from Hypsilurus such as hair-like skin receptors in Gonocephalus or lens-like skin receptors in
Hypsilurus (Ananjeva & Matveyeva-Dujsebayeva 1996), differing karyotypes (Ota et al. 1992) and differences in
hemipenis morphology (Böhme 1988).
The first Gonocephalus species to be discovered was Gonocephalus chamaeleontinus (Laurenti, 1768),
originally described as Iguana chamaeleontina. The genus Gonocephalus was erected much later by Kaup (1825:
590) with Lophyrus tigrinus Duméril & Bibron, 1837 (=G. chamaeleontinus) constituting the type species (fide
Wermuth 1967). The original name was Gonocephalus (gono- from gr. γωνία / gonia - angle, - cephalus from gr.
κεφαλή / kephale - head) but in a later publication (Kaup 1827: 614) he amended it to Goniocephalus. Wagler
(1830: 150) introduced the alternative spelling Gonyocephalus that was used by several authors in subsequent
publications (e. g. Gray 1845, Boulenger 1885, 1908, 1912, Smith 1922).
Since the description of Gonocephalus chamaeleontinus the diagnostic characters changed over time in order
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to accommodate new species. The initial characteristic of an angular head was mainly based on a well developed
Canthus rostralis and a strongly raised superciliary edge which nearly forms a triangle in G. chamaeleontinus.
Apart from a transverse gular fold and enlarged scales on either side of the dorsal and nuchal crest several other
characters are mirrored in many Asian agamid lizards. Through including species such as G. bellii (Duméril &
Bibron, 1837) and G. liogaster (Günther, 1872) or Philippine Gonocephalus, i.e. G. sophiae (Gray, 1845), G.
interruptus Boulenger, 1885 and G. semperi (Peters, 1867), and species of the Australian radiation (nowadays
Hypsilurus) the diagnostic character of a strongly raised angular superciliary edge had to disappear as most of these
species show rather rounded and not angular canthi.
Boulenger (1885) summarised the genus diagnosis as follows: “Tympanum distinct. Body compressed. Dorsal
scales small, uniform or intermixed with larger ones. A dorsal crest. A strong transverse gular fold. Male with a
gular sac. No preanal or femoral pores.” This diagnosis referred to currently three accepted genera, i. e.
Gonocephalus, Hypsilurus and Coryphophylax.
Several years later Boulenger (1908) described Gonyocephalus [sic] robinsonii, a mountainous form of agamid
lizard from Gunung (=Mt.) Tahan (Pahang, West Malaysia). Having only a single specimen at hand with no other
known agamid lizards showing a comparable set of characters he assigned it to the genus Gonocephalus where at
least some resemblance could be found and characters were mirrored. However, at the end of his original
Gonocephalus robinsonii publication Boulenger (l.c.) wrote: “A remarkable form, unlike anything previously
described”.
In the following we will endeavour to evaluate the taxonomic status of Gonocephalus robinsonii and whether it
is plausible or appropriate to partition the genus into several genera.
Material and methods
Meristic and morphometric data were recorded from type specimens and additional material where species
association is unambiguous. Measurements were taken using a sliding calliper with a precision of 0.1 mm or using
a ruler with a precision of 1 mm (SVL: snout–vent length; TL: tail length; HL: head length; HW: head width).
Additionally, photographic material of living specimens collected during field trips was available for further
comparisons. For our morphometric studies we examined a total of close to 200 specimens from all currently
known species (with the exception of the Philippine G. interruptus) of the genus Gonocephalus distributed over all
age classes where available. A complete list including sex and locality for each specimen would lengthen Appendix
1 tremendously. Hence we decided to present these data for some representative specimens that are either type
specimens or individuals that show all the characteristics of a particular species. For the remaining material only
their collection number is given. The museum material used for morphological comparisons is listed in Appendix
1. Abbreviations: BMNH (British Museum of Natural History London); FMNH (Field Museum of Natural
History); MNHP (Muséum National d'Histoire Naturelle Paris); MTKD (Museum für Tierkunde Dresden); RMNH
(Rijksmuseum van Natuurlijke Historie, now Naturalis Biodiversity Centre; merged with ZMA); SMF
(Senckenberg Forschungsinstitut und Naturmuseum); USNM (National Collection United States National Museum
of Natural History, Smithsonian Institution); ZFMK (Zoologisches Forschungsmuseum Alexander Koenig); ZMA
(Zoölogisch Museum Amsterdam, now Naturalis Biodiversity Centre; merged with RMNH); ZMB (Zoologisches
Museum Berlin, Museum für Naturkunde); ZRC (Zoological Reference Collection, Singapore).
Micro-tomographic analysis was performed at the Museum für Naturkunde Berlin using a Phoenix nanotom
X-ray|s tube at 80–90 kV and 130–200 µA, generating 1000–1440 projections per scan with 750–1000 ms.
Differences in beam- and projection-settings depended on the respective specimen size. Effective voxel size ranged
between 14–21 µm. The cone beam reconstruction was performed using the datos|x-reconstruction software (GE
Sensing & Inspection Technologies GmbH phoenix|x-ray) and the data were visualized and analysed in VG Studio
Max 2.1.
Genetic data for 16S ribosomal RNA of G. robinsonii were retrieved from the Nucleotide database (GenBank)
of the National Centre for Biotechnology Information (www.ncbi.nlm.nih.gov/nuccore). Subsequently we searched
for most similar sequences using the Basic Local Alignment Search Tool (BLASTN 2.2.28+; megablast
subroutine) (Zhang et al. 2000). The resulting data set was searched for species belonging to the agamid subfamily
Draconinae. A neighbour joining tree representation was produced using a subroutine of the same program
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(BLAST Tree View) that computes pairwise alignment of the preselected sequences. Several species had multiple
data sets and only those were included that had been used in previous studies to allow for comparisons.
Subsequently we applied a more stringent Fast Fourier Transform method (Katoh & Standley 2013; MAFFT, Q-
INS-I refinement) for the alignment to compare the resulting tree to that obtained from BLAST. In the final step we
employed the program SeaView 4.5.4 (Gouy et al. 2010) to evaluate exclusively the phylogenetic relationships
within the draconine clade. This program uses Clustal Omega (Sievers et al. 2011) for the alignment procedure as
well as PHYLIP 3.696 / dnapars (Felsenstein 1989) and PhyML 3.1 (Guindon et al. 2010) to calculate
parsimonious and maximum likelihood phylogenetic trees, respectively. In both cases the number of bootstrap
replicates was set to 100 (non parametric bootstrap analysis of the best tree). In PhyML the best tree was found by
enabling both nearest neighbour interchange (NNI) and subtree pruning and regrafting (SPR). The starting tree for
the maximum likelihood diagram calculation was optimized for tree topology by adding five random starts.
Genetic distances were calculated using the Jukes-Cantor (J-C) model (http://www.insilicase.co.uk/Web/
JukesCantor.aspx; Jukes & Cantor, 1969) based on the results from PhyML statistics report (uncorrected genetic
distance, no. of complete sites, no. of variable sites). The number of complete sites taken into account for
alignment varied from 396 to 402. GenBank accession numbers of the material used here and within the respective
previous phylogenetic studies by Honda et al. (2002) and Pyron et al. (2013) are listed in Appendix 2.
Results
Morphology. In their short review of the genus Gonocephalus Manthey & Denzer (1991, 1992a & b, 1993)
proposed species groups based on morphological similarities. These comprised the Gonocephalus bellii/bornensis
Group (G. bellii, G. beyschlagi Boettger, 1892, G. bornensis (Schlegel, 1848) [including G. denzeri Manthey, 1991],
G. liogaster and G. semperi, G. sophiae, G. interruptus from the Philippines [included later by Denzer & Manthey
2009]), the Gonocephalus chamaeleontinus Group (G. chamaeleontinus, G. kuhlii (Schlegel, 1848), G. doriae
Peters, 1871 and G. abbotti Cochran, 1922) and the Gonocephalus megalepis Group (G. klossi Boulenger, 1920, G.
megalepis (Bleeker, 1860) and G. lacunosus Manthey & Denzer, 1991). Gonocephalus grandis, G. m jo be rg i and G.
robinsonii were not assigned to any of these groups. A later study (Denzer & Manthey 2009) of the type specimen
of G. mjobergi revealed that this species does most certainly not belong to the genus and it was therefore suggested
that it should be considered as incertae sedis and rather be called Genus A within Gonocephalus sensu lato. In the
same paper Denzer & Manthey (l. c.) pointed out some morphological similarities between G. mjobergi and
Ptyctolaemus Peters, 1864 as well as Mantheyus Ananjeva & Stuart, 2001.
G. robinsonii exhibits morphological and meristic characters that distinguish it from all other Gonocephalus
species, in particular from the type species of the genus, i. e. Gonocephalus chamaeleontinus. One of the diagnostic
characters of the genus is the presence of a transverse gular fold. However, G. robinsonii does not possess such a
fold but rather an antehumeral fold that is extending towards the sides of the neck. The gular sac is very large
reaching onto the chest—hence preventing the development of a transverse gular fold—unlike most other species
of Gonocephalus (an exception being G. megalepis from Sumatra). From our field work and observations in
captivity we learned that G. robinsonii can extend and fold its gular sac in a rapid sequence similar to what has been
described as display behaviour of Draco species. Such behaviour has not been observed in any other
Gonocephalus. Additionally the shape of the gular sac in G. robinsonii is different to that of all other species of
Gonocephalus as it has a long narrow rounded tip (Figure 1).
In all Gonocephalus species but G. robinsonii the nuchal (and sometimes the dorsal) crest is separated laterally
from the nuchal scales (or dorsal scales, respectively) by one or more rows of enlarged scales. In G. robinsonii the
nuchal crests is formed by individual triangular scales with no significantly enlarged scales on either side, as is true
for the dorsal crest.
Morphometrically Gonocephalus robinsonii differs from G. chamaeleontinus Group in its head and body
proportions. While TL/SVL = 1.49–1.87 in G. chamaeleontinus (n=27), 1.33–1.54 in G. doriae (n=9), in G. abbotti
1.38–1.64 (n=7) and 1.7–2.07 in G. kuhlii (n=26) this ratio comes to 2.27–2.60 (n=4) in G. robinsonii reflecting the
much longer tail of the latter. The elongated head shape of G. robinsonii is represented by the HW/HL ratio of
0.54–0.62 whereas that ratio equates to 0.62–0.75 in G. chamaeleontinus, 0.62–0.69 in G. d o r i a e , 0.58–0.69 in G.
abbotti and 0.63–0.75 in G. kuhlii. Only G. grandis has a similarly pointed head shape with a HW/HL ratio of 0.52–
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0.60 while all other species have HW/HL ratios in the region between 0.6–0.8. However, G. robinsonii does not
show particular morphometric differences to all Gonocephalus species. With respect to its TL/SVL proportions it
falls within the range of several species such as G. bellii (TL/SVL 2.2–2.36, n=14) or G. semperi (2.04–2.28, n=2).
At its upper end the ratio of TL/SVL (2.27–2.60, n=5) is one of the highest as compared to other species of this
genus. It is only exceeded by inc. sed. Genus A (Gonocephalus) mjobergi (TL/SVL= 3.01, n=1) as well as G.
grandis (TL/SVL=2.60–2.80, n=6) and matched by G. beyschlagi (TL/SVL=2.16–2.55, n=13), G. bornensis (TL/
SVL= 1.95–2.64, n=52 incl. types of G. denzeri) and G. liogaster (TL/SVL=2.14–2.46, n=12).
FIGURE 1. Ventrolateral view of the head, gular and pectoral regions (“Gonocephalus robinsonii holotype BMNH
1946.8.14.81)
Furthermore, Gonocephalus robinsonii has a pair of distinctive occipital protuberances that are not observed in
any other species of Gonocephalus. In living and preserved specimens this structure is most prominent close to the
posterior supraocular region and subsequently decreasing in height to a less prominent ridge that runs towards the
onset of the nuchal crest. From a dorsal view in live the protuberances appear V- or U-shaped with the onset of the
nuchal crest forming the centre. The ridges are covered with slightly enlarged scales as compared to the
surrounding ones. CT scans reveal that this structure is part of the skull morphology and consists of bony
protrusion of the parietal (Figure 2). These protrusions run along the outer edge of the parietal and are connected by
a straight and lower rim halfway behind the frontoparietal foramen and the posterior end of the parietal. Among
agamid lizards of the subfamily Draconinae prominent occipital bony ridges have otherwise only been described
by Manthey & Grossmann (1997) for the Sumatran monotypic genus Dendragama, i. e. Dendragama boulengeri
Doria, 1888.
The cranial skeleton of Dendragama boulengeri (see Figure 2B) clearly differs from that of Gonocephalus
robinsonii (see Figure 2A). In particular the form and shape of the parietal shows significant differences. While the
parietal is nearly semicircular in D. boulengeri this bone is rather formed like a V with a broadened base in G.
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robinsonii. Additionally the surface and prominent lateral ridges of the parietal bone are rugose in D. boulengeri
but near smooth in G. robinsonii. In D. boulengeri the nasals are merely in contact as the tip of the frontal extends
close to the premaxilla. In G. robinsonii the nasals are in broad median contact with the tip of the frontal hardly
only extending so that the posterior 30% of the nasals are not in contact. Again the surface structure of nasals and
premaxilla in D. boulengeri is rather rugose with some conical prominences while that of G. robinsonii is nearly
smooth.
FIGURE 2. A. Dorsal view of the skull of Gonocephalus robinsonii (ZMB 48856); B. Dorsal view of the skull of
Dendragama boulengeri (ZMB 54503); C. Lateral view of the skull of “ Gonocephalus robinsonii showing the elevated edges
on the parietal (ZMB 48856); D. Posterior view of the skull of “ Gonocephalus robinsonii with the parietal bone coloured
green (ZMB 48856). Scale bar = 4mm.
We would also like to note that species of the Gonocephalus chamaeleontinus Group possess a well defined
supraorbital arch formed by a forward curved procession of the frontal and a large backward curved procession of
the prefrontal (Plate 23 Figure 3 in Schlegel [1837] for Galeotes lophyrus = Gonocephalus chamaeleontinus or
Gonocephalus kuhlii; ambiguous identification). As can be seen in Figures 2A, 2C & 2D G. robinsonii does not
possess a supraorbital arch but only shows small spines at the posterior edge of the frontal and prefrontals,
respectively. In Dendragama boulengeri the prefrontal has a well developed backward pointing procession (Figure
2B) but no forward directed procession of the frontal or postorbital and hence it is missing the posterior part of a
supraorbital arch.
Karyotype. Chromosomal investigations of five Gonocephalus species by Diong et al. (2000) yielded two
different karyotypes within the genus with all but G. robinsonii 12M + 20m (2n=32) having a karyotype of 22M +
20m (2n=42). The latter karyotype was present in G. chamaeleontinus, G. liogaster, G. bellii, G. grandis (specimens
from peninsular Malaysia and Borneo) and G. miotympanum [=G. bornensis fide Manthey & Denzer 1992b]).
Diong et al. (l.c.) concluded that including G. robinsonii in Gonocephalus would render the genus paraphyletic.
However, they did not suggest removing G. robinsonii from the genus without further studies but they hypothesized
that G. robinsonii could be related to Australian agamids where this karyotype is common, e.g. Chlamydosaurus
kingii or Pogona vitticeps.
Later Ota et al. (2002) revisited the initial chromosomal study by Diong et al. (l.c.) and found that the same
kind of karyotype revealed by G. robinsonii is shared with Acanthosaura armata (Hardwicke & Gray, 1827) and
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certain populations of Calotes versicolor (Daudin, 1802). They concluded that G. robinsonii is clearly not a
member of the Australian agamid radiation but belongs to arboreal radiation of Southeast Asian agamid lizards that
are nowadays collectively considered as the subfamily Draconinae.
It should be noted that karyotyping alone cannot serve as a taxonomic character and may only give an
indication for speciation. The number of chromosomes can even differ between populations within a single species
as has been shown by Ota (1988) in the case of Japlura swinhonis Günther, 1864 where three different sets were
found (2n=46, 40 and 36 chromosomes, respectively).
Molecular phylogenetics. A phylogenetic study of Honda et al. (2002) using mitochondrial 12S and 16S
rRNA nucleotide material included G. chamaeleontinus, G. grandis, G. miotympanum (= bornensis fide Manthey &
Denzer 1992b) and G. robinsonii led to the result that the 2n=42 karyotype group of Gonocephalus appears to be
monophyletic while G. robinsonii turns out to be more closely related to Japalura polygonata (Hallowell, 1861)
(2n=46) within a node that additionally comprises Acanthosaura crucigera Boulenger, 1885. From a study by
Guha & Kashyap (2006) related to a case of food poisoning and using forensically informative nucleotide
sequencing (16S rRNA) one can draw the conclusion that G. robinsonii is more closely related to Japalura
polygonata than to Calotes versicolor or Aphaniotis fusca Peters, 1864. Other species of the genus Gonocephalus
were unfortunately not included but still the result corroborates that of Honda et al. (l.c.). Pyron et al. (2013) used a
supermatrix approach to evaluate phylogenetic relationships of 4161 squamate species based on currently available
sequence data. With respect to the genus Gonocephalus their study included specimens of G. chamaeleontinus, G.
grandis, G. kuhlii and G. robinsonii. While the first three species turned out to form a monophyletic group G.
robinsonii again clustered with Japalura polygonata. The authors considered both genera Gonocephalus and
Japalura in their current composition as non-monophyletic.
We conducted an alignment test on GenBank using 16S ribosomal RNA sequences that included the available
data sets for Gonocephalus species and those of other species of the subfamily Draconinae from Southeast Asia.
The routine returned 25 Southeast Asian species from the subfamily Draconinae. As the genus Draco Linnaeus,
1758 was overrepresented with a total of eleven species only two species were included into the initial alignment
study in order to keep the resulting tree less congested and clearer. When all Draco species were included into the
simulation they came out as one monophyletic group. The sequence of an unidentified Pseudocalotes sp.
(EU502979) was disregarded but Japalura luei Ota, Chen & Shang, 1998 from Taiwan was included to facilitate an
intrageneric comparison with Japalura polygonata (from Japan). Of the remaining species (n=16) those with the
highest sequence matching score—in cases where more than one sequence was available—were used to construct a
neighbour joining diagram. Most of the material has been used in previous studies by Honda et al. (in part, 2000 &
2002) and by Pyron et al. (l.c.), respectively (see Appendix 2). As outgroups we chose Hypsilurus “godeffroyi”
Peters, 1867 a Melanesian species formerly assigned to Gonocephalus, Physignathus cocincinus Cuvier, 1829 an
Indochinese species currently considered to be a sister taxon or member of the Australian-Melanesian subfamily
Amphibolurinae (as are all Hypsilurus species) and Hydrosaurus amboinensis (Schlosser, 1768) a species from
Wallacea representing the recently erected subfamily Hydrosaurinae. As the identity of Hypsilurus "godeffroyi"
(GenBank AB031984; KUZ 45215, Kyoto University, Dept. of Zoology, collected in Irian Jaya) on species level is
questionable -this species is not known to occur on New Guinea- we place the species epithet within quotation
marks. The results of 16S sequence alignments by BLAST are depicted in Figure 3. The resulting NJ trees from
BLAST and MAFFT were near identical with the only major exception that Gonocephalus bornensis forms a clade
with Bronchocela cristatella when MAFFT is used (s. below for maximum likelihood and most parsimonious
trees).
As can be seen in Figure 3 this comparatively straightforward method recovers previously published
relationships such as the position of Physignathus in close relationship to the subfamily Amphibolurinae as well as
the subfamilial status of Hydrosaurinae (Honda et al. 2000 & 2002 Schulte et al. 2003, Pyron et al. 2013). Within
the Draconinae monophyly of Draco and Acanthosaura are established as is the isolated position of Mantheyus.
Most importantly the neighbour joining dendrogram clearly shows that G. robinsonii resides outside the 2n=42
karyotype group of Gonocephalus species that forms its own clade. It reveals that Gonocephalus becomes
paraphyletic if G. robinsonii remains a member of this genus. From the diagram it can be concluded that G.
robinsonii is more closely related to Japalura polygonata and J. luei than to any member of the genus
Gonocephalus corroborating the results published by Honda et al. (2002, Fig. 1B; they did not include J. luei). In
this respect our tree is also consistent with the results presented by Pyron et al. (2013). Their study showed that
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additionally G. k u h l ii —s. below for comments on the identity of this specimen—belongs to the clade consisting of
all Gonocephalus species but G. robinsonii. Combining these results it appears plausible to form a phyletic group
containing G. chamaeleontinus, G. grandis, G. kuhlii and G. miotympanum (=bornensis) that could be considered as
Gonocephalus sensu lato excluding G. robinsonii and G. (Genus A) mjobergi) as the genus would otherwise be
paraphyletic. If karyotype results (see above) are taken into account Gonocephalus s. l. also includes G. bellii and
G. liogaster—two species with a karyotype of 2n=42 and morphologically very similar to G. bornensis.
FIGURE 3. Neighbour joining dendrogram of the subfamily Draconinae with outgroups represented by Hypsilurus,
Physignathus (Amphibolurinae) and Hydrosaurus (Hydrosaurinae) resulting from BLAST based on 16S ribosomal RNA
sequence alignments.
Subsequently we reduced our analysis to species belonging to the subfamily Draconinae. For clarity the genera
Japalura, Acanthosaura and Draco are only represented by a single species (total no. of species included n=13).
The alignment procedure using Clustal Omega using the same sequences as in BLAST / MAFFT found 383
complete sites of which 160 were variable and 104 informative. Figure 4a depicts the maximum likelihood (ML)
diagram (NNI / -ln[L]= 2686.8) for the draconine lizards resulting from our PhyML analysis. Figure 4b shows the
most parsimonious (MP) phylogenetic tree computed using the dnapars routine from PHYLIP. In both cases G.
robinsonii and Japalura polygonata form a clade well separated from the other Gonocephalus species.
Additionally a closer relationship between G. bornensis and Bronchocela cristatella than to the other two
Gonocephalus species is supported as has already been seen in our MAFFT analysis. The major difference between
BLAST/MAFFT analysis and the maximum likelihood / most parsimonious diagrams can be seen in the position of
Calotes versicolor which no longer clusters with G. robinsonii and Japalura polygonata. The main difference
between ML and MP trees is the position of Acanthosaura crucigera. However, it should be noted that the branch
support for a clade including A. crucigera and Phoxophrys nigrilabris (ML tree) is considerably higher than
placing this species within the Gonocephalus / Bronchocela clade (MP tree).
The genetic distances resulting from the analysis clearly support our hypothesis that G. robinsonii is only
remotely related to other Gonocephalus species. Within the Gonocephalus species group the closest relationship
was found to exist between G. chamaeleontinus and G. grandis (uncorrected genetic distance 7.2%, J-C 7.58%). G.
robinsonii J-C genetic distances were calculated to be 13.24%, 16.02% and 22.92% for G. chamaeleontinus, G.
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grandis and G. bornensis, respectively. The latter value (G. robinsoniiG. bornensis) nearly equals the distance
between G. robinsonii and Mantheyus phuwuanensis (J-C 23.47%) or Draco volans (J-C 22.51%). These values
clearly show that G. robinsonii is not closely related to any of the other studied species of the genus Gonocephalus.
None of our tree topologies supports monophyly of the genus Gonocephalus. Both PhyML and PHYLIP analysis
also support the close relationship between G. chamaeleontinus and G. grandis. Additionally the large genetic
distances between G. bornensis and G. chamaeleontinus as well as G. grandis (J-C 14.54% and 16.69%,
respectively) indicate that even if G. robinsonii is removed from Gonocephalus the genus is genetically highly
diverse and probably still paraphyletic.
FIGURE 4. A. Most parsimonious phylogenetic tree resulting from PHYLIP/Dnapars analysis. B. Maximum likelihood
phylogenetic tree resulting from PhyML analysis Branch length scale represents number of substitutions/site. Branch support
values are given above the branch. The trees were re-rooted for Mantheyus phuwuanensis.
When we tried to corroborate the phylogenetic result of Pyron et al. (2013) and our previous morphological
study (Manthey & Denzer 1993) with respect to a close relationship between G. chamaeleontinus and G. k u h l i i by
using a different set of sequences (C-mos as nuclear marker) we found that their assignments in GenBank may be
incorrect. The identification of either of the sequences for G. kuhlii (GenBank DQ 340681) or for G.
chamaeleontinus (GenBank AF137526) is probably erroneous. The sequence alignment (using Clustal Omega for
deposited c-mos sequences) resulted in a genetic difference of 0.1% (transition-transversion ratio close to 0 with
only 1 out of 923 sites variable). With such a small genetic difference these two specimens have to be conspecific.
According to Hugall et al. (l. c.) the respective specimen (AMS R126130) for DQ340681 (= G. kuhlii, see above)
was collected in Cibodas Forest, Java, Indonesia which would coincide with the distribution of G. k u h l i i (and
potentially G. chamaeleontinus). Locality data for the specimen corresponding to G. chamaeleontinus (GenBank
AF137526) are neither given on GenBank nor are we aware of a publication that may have provided further details.
For the time being we prefer to abstain from a further analysis of this particular subset within Gonocephalus until
specimens and associated gene material have been identified unambiguously.
Discussion
On the basis of chromosomal and molecular investigations in conjunction with morphological and meristic data we
suggest all currently recognised species remain included in the genus Gonocephalus with the exception of G.
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robinsonii and G. (Genus A) mjobergi. Members of the chamaeleontinus Group (G. chamaeleontinus, G. d o ri ae , G.
kuhlii and G. abbotti [= G. doriae abbotti fide Manthey & Denzer 1993]) are the closest relatives to the type species
of the genus and we consider this group as Gonocephalus s. str. The remaining species groups (megalepis Group,
bellii / bornensis Group and G. grandis) we consider as a clade Gonocephalus s. l. For G. robinsonii we propose to
remove it from the synonymy with Gonocephalus as the genus would otherwise constitute a paraphyletic clade.
From the phylogenetic analysis it would seem appropriate to include Gonocephalus robinsonii into the genus
Japalura. However, Japalura spp. are morphologically dissimilar (e. g. small or no gular sac, dorsal scales
heterogeneous) and do not even share a distribution in the same zoogeographical subregion. The closest locality
record of any Japalura species would be that of Japalura cf. yunnanensis in northwest Thailand (Manthey &
Denzer 2012). Additionally the genus Japalura turned out to be a highly paraphyletic group itself (Pyron et al. l.c.)
and transferring G. robinsonii to Japalura would add even more to the current confusion. Additionally, the JC
distance between G. robinsonii and J. polygonata turns out to be 19.17%.
In our BLAST dendrogram Calotes versicolor appears in the same node as Gonocephalus robinsonii /
Japalura species implying a possible relationship. The genus Calotes has been shown to be monophyletic by Pyron
et al. (l. c.; 11 species included) and inclusion of G. robinsonii into this genus would render it paraphyletic. The
pairwise JC genetic distance between G. robinsonii and C. versicolor amounts to 12.74%. All species of Calotes are
additionally morphologically different from G. robinsonii. They possess keeled dorsal scales that are typically
equal or larger than the ventral scales. The gular pouch is much smaller than that of G. robinsonii and its scales are
enlarged in most species.
Another genus apparently related to Gonocephalus robinsonii based on our BLAST/MAFFT analysis is
Acanthosaura. Again G. robinsonii is morphologically dissimilar and does not share any of the genus characters of
Acanthosaura, e.g. postorbital spine, interrupted nuchal and dorsal crests or shape of crest scales. Inclusion of G.
robinsonii into Acanthosaura would result in paraphyly of this otherwise monophyletic group. Additionally
Phoxophrys nigrilabris (Peters, 1864) shares the node with Acanthosaura. However, all currently recognized
members of Phoxophrys are small lizards (SVL max. 85 mm) inhabiting Borneo and Sumatra sharing several
features such as a concealed tympanum and a blue inside of the mouth. G. robinsonii does not share any of these
characters and is additionally differing by its large gular pouch. Genetically G. robinsonii and A. crucigera as well
as P. nigrilabris have a JC distance of 15.41% and 18.42%, respectively.
Morphologically Gonocephalus robinsonii shares several characters with Dendragama boulengeri, a
monotypic genus endemic to Sumatra. This includes the bony ridges in the occipital region as well as
morphometric data (TL/SVL 2.1–2.4) and meristic characters (nuchal and dorsal crest formed by individual
scales). However, D. boulengeri differs from G. robinsonii in a set of morphological features that define its generic
status. In D. boulengeri the nuchal crest sits on top of a nuchal sail. The crest scales do not have the flat triangular
appearance as in G. robinsonii but are rather of a spine-like shape. D. boulengeri possesses conical scales on the
posterior part of the hindlimbs sometimes present but hardly discernible in G. robinsonii). The gular sac is
comparatively small and not shaped as in G. robinsonii (large with a rounded tip). Additionally as mentioned before
both species differ in several characters with respect to the form and shape of cranial skeletal elements (see Figure
2).
Conclusion. Albeit that erecting yet another monotypic genus does not contribute much to the knowledge of
the phylogenetic relationships among draconine lizards we are confident that—on the grounds of the evidence
compiled—taking the taxonomic decision of erecting a new genus to accommodate Gonocephalus robinsonii is
justified.
Malayodracon gen. nov.
Type species: Gonyocephalus robinsonii Boulenger, 1908; Type locality: Gunong (=Gunung / Mt.) Tahan (5,200 ft
/ 1,585 m), Pahang, Malaysia; the holotype (by monotypy) is deposited in the British Museum Natural History
BMNH 1946.8.14.81 (previously BMNH 1906.2.28.8); collected by H.C. Robinson.
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Taxon—Malayodracon robinsonii (Boulenger, 1908)
Chresonymy
Gonyocephalus robinsonii—Boulenger 1908: 65; Boulenger 1912: 67
Gonyocephalus robinsoni—Smith 1922: 269
Gonocephalus robinsoni—Smith 1930: 24; Smirnova 2003: 128
Goniocephalus robinsoni—Smith 1935: 133; Sly 1976: 156; Bourret 1947 “2009”: 211
Gonocephalus robinsonii—Smedley 1931: 110; Wermuth 1967: 61; Moody 1980: 299; Denzer & Manthey 1991: 312;
Manthey & Denzer 1992a: 16; Manthey & Schuster 1992: 65; Manthey & Grossmann 1997: 189; Chan-ard et al. 1999:
102; Diong et al. 2000: 73; Honda et al. 2002; Denzer & Manthey 2009: 256; Grismer 2011: 258; Pyron et al. 2013
(Gonocephalus inc. sed.) robinsonii—Manthey 2010: 46
Diagnosis. Medium sized lizard; males: SVL 115–158 mm, TL 285–320 mm; females: SVL 115–132 mm, TL
290–300 mm (type specimen, male, SVL 152 mm, TL 320 mm), TL up to 2.6 times SVL; body triangular in cross-
section; no patagia; head large, elongated and pointed; bony protuberances on the occipital region; sharp canthus
rostralis and rounded superciliary edge; tympanum exposed; antehumeral fold present, transverse gular fold absent;
large gular sac; nuchal and dorsal crest continuous; no preanal or femoral pores.
Comparison to other draconine genera. Morphologically Malayodracon robinsonii differs from all other
agamid lizards by a combination of the following characters: visible tympanum, bony ridge in the occipital region,
large gular sac with a rounded tip extending onto the chest, small weakly keeled, muricate dorsal scales and
triangular crest scales.
From its former congeners of the genus of Gonocephalus s. l. it differs additionally by the absence of
supporting scales along the nuchal crest (vs. present) and the absence of a distinct transverse gular fold (vs.
present). In particular Malayodracon robinsonii differs from the Gonocephalus bellii Group by the possession of
triangular nuchal and dorsal crest scales (vs. spines), from the G. megalepis Group by the lack of a large conical
scale at the posterior joint of mandibular and maxillar region (vs. present in lacunosus, megalepis and klossi), from
the G. chamaeleontinus Group by the shape of the supraciliary edge (rounded in M. robinsonii vs. angular) and the
form of the nuchal crest (single triangular scales in M. robinsonii vs. erected nuchal crest consisting of several scale
rows) and G. grandis by the shape of the nuchal crest (crest scales of G. grandis fused similar to a sail).
Malayodracon robinsonii differs from the species that appear in the same node of our phylogenetic analysis as
follows: from all Acanthosaura by a lack of a large spine behind the orbit (vs. present in Acanthosaura) and from
all Phoxophrys by an exposed tympanum and the absence of a blue mouth. From Japalura polygonata and J. luei it
can be distinguished by the possession of small dorsal scales (vs. large heterogeneous scalation) as well as the
possession of a dorsal crest. From all other species of Japalura it can be distinguished by its gular sac pholidosis
which is small and smooth in M. robinsonii and enlarged and often keeled in Japalura species. Additionally most
Japalura spp. have a concealed tympanum (exceptions are some Indian and Nepalese species formerly referred to
as Oriotiaris (Kästle & Schleich 1998). From Calotes versicolor and all other species of that genus M. robinsonii
differs in pholidotic characters such as small dorsal and gular scales (vs enlarged dorsals in C. versicolor) and
dorsal scales smaller than ventrals (vs. dorsal scales larger than ventral scales in C. versicolor) as well as the shape
of the nuchal and dorsal crest.
Furthermore it differs from the remaining Southeast Asian genera in the subfamily Draconinae as follows:
Aphaniotis Peters, 1864 and Pseudocophotis Manthey in Manthey & Grossmann, 1997 by a visible tympanum;
additionally by absence of a blue mouth or a rostral appendage and from Pseudocophotis by the absence of a
prehensile tail
Bronchocela Kaup, 1827 by the absence of a lateral skin fold on both sides of the neck
Complicitus Manthey in Manthey & Grossmann, 1997 by the absence of lateral pockets on the gular pouch
Coryphophylax Fitzinger in Steindachner, 1867 by the absence of a nuchal sail and the possession of a dorsal
crest
Dendragama Doria, 1888 by cranioskeletal features and gular pouch size
Draco Linnaeus, 1758 by the absence of elongated ribs and patagia
Harpesaurus Boulenger, 1885 and Thaumatorhynchus Parker, 1924 by the absence of a rostral appendage
Hypsicalotes Manthey & Denzer, 2000 by the absence of large plates on either side of the head and the absence
of large lanceolate scales along the midline of the gular pouch
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Mantheyus Ananjeva & Stuart, 2001, Ptyctolaemus Peters, 1864 and “Gonocephalus” (Genus A) mjobergi
Smith, 1925 by the absence of longitudinal skin folds on the gular region
Pseudocalotes Kaup, 1827 by the absence of rhombic dorsal scales
General description. The holotype (see Figure 1) has been described in Boulenger (1908) and several aspects
including variation were added by Smedley (1931) after additional material became available. A very detailed
description of Malayodracon robinsonii (as Gonocephalus robinsonii) is given in Grismer (2011).
FIGURE 5. Malayodracon robinsonii from Tanah Rata, Cameron Highlands, West Malaysia. Photo: U. Manthey.
Head scales small; scales on the supraciliary edge longer than wide; nasal in contact with first supralabial,
sometimes also with second; typically 8 supra- and 8 infralabial scales (maximum 10 each). Slightly enlarged
scales on the prominent v-shaped protrusion in the occipital region; large conical scale on either side of occiput.
Gular pouch large, reaching the chest, tip rounded; gular scales small, homogeneous, smooth or slightly keeled.
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Body elongated, laterally slightly compressed; dorsal scales nearly homogeneous, small, keeled, directed
backwards and upwards intermixed with several—usually five or six—parallel oblique rows of enlarged scales.
Scale numbers around midbody sometimes exceeding 100. Nuchal and dorsal crests continuous consisting of a
single row of flat triangular—in some specimens lanceolate—scales touching at the base and decreasing in size
caudad. Ventrals significantly larger than dorsals, keeled. Subdigital lamellae keeled, 31–36 on fourth toe. Tail
long, roundish in cross-section. Enlarged scales along the mid ridge of the tail, continuous with dorsal crest.
Underside of tail with strongly keeled scales.
Adults in life typically dark green dorsally and dorsolaterally, dirty white ventrally; sometimes ground colour
yellowish green (see Figure 5). Gular sac typically reddish brown, greyish or black, continuously darkening
towards the tip. A dark shoulder patch and dark oblique cross bands often visible. Enlarged dorsal scales (if
present) often coloured much lighter than surrounding body colouration, forming bands. Labials and area below
tympanum (plus tympanum) conspicuously white. Colour dependent upon age. Juveniles typically yellowish
brown ground colour with brown cross bands. Labials in juveniles rather dirty white to pale brown, shoulder patch
brown. In alcohol the colour fades and grey tones are prevalent (s. photograph of the type specimen in Figure 1).
However, the black shoulder patch and dark dorsal cross bands remain discernible.
Va ri at io n . Hitherto known specimens from the type locality (Gunung Tahan) do not show enlarged dorsal
scales arranged in oblique rows (Boulenger 1908, Sly 1976) as it can be seen in specimens from the Cameron
Highland region. It is conceivable that these two populations have been separated for a long time and constitute
subspecies. However, in order to establish consistency of this character more material from the remote mountain
ranges of central Malaysia is needed. Additionally there exists a photographic record of a specimen from the
Cameron Highlands without apparent enlarged scales across the dorsum rendering the above observation doubtful.
Distribution. Malayodracon robinsonii is restricted in its distribution from mid to high altitude areas of the
peninsular Malaysian highlands (approx. 600–1500 m asl) inhabiting submontane and montane forests. M.
robinsonii shares this isolated distribution with several other reptile species that are restricted to the Cameron
Highlands or the central mountain ranges of Malaysia (Fraser and Larut Hills; Genting Highlands) such as
Trimeresurus nebularis Vogel, David & Pauwels, 2004, Hebius sanguineus (Smedley, 1931), Macrocalamus
tweediei Lim, 1963 and Collorhabdium williamsoni Smedley, 1931. Endemism on genus and species level is
comparatively high in this geographical area.
Malayodracon robinsonii has been reported from Genting Highlands (Ulu Kali), Cameron Highlands (Tanah
Rata, Gunung Brinchang, Gunung Beremban, Gunung Jesar) and further north from its type locality Gunung
Tahan.
Etymology. The name Malayodracon was chosen to express that the type and currently only known species of
the genus is restricted to Malaysia (latinized malaya, male form malayo owing to the gender of the ending -dracon)
and constitutes a genus belonging to the subfamily Draconinae (gr. drakon / δράκων; a serpent in Greek mythology;
latinized dracon = engl. dragon).
Acknowledgements
We would like to thank C. McCarthy (London) for giving us access to the type specimen of Gonyocephalus
robinsonii. We appreciate the help of M. S. Hoogmoed (Leiden), F. J. Obst (Dresden), G. Köhler (Frankfurt), A.
Resetar (Chicago), R. Günther, M.-O. Rödel & F. Tillack (Berlin), R. Gemel & H. Grillitsch (Wien) and K. Lim
(Singapore) as well as curatorial staff of the herpetological collections in Amsterdam and Paris for facilitating the
examination of specimens in their care.
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APPENDIX 1. Material examined.
Gonocephalus (= Malayodracon) robinsonii BMNH 1946.8.14.81 (type; adult male; Gunung Tahan, 5200 ft); ZMB 48878
female, ZMB 48848 semiadult, all from Tanah Rata, Cameron Highlands, Malaysia; ZMB 48856 (one adult male, one
adult female from Gunung Jesar, Cameron Highlands, Malaysia)
Gonocephalus abbotti ZFMK 41710 & 44274 (two males, Malaysia); ZMB 48924 (female, West Malaysia);
Gonocephalus bellii ZFMK 51162 (male, Pulau Pinang, Malaysia); ZFMK 21493 & 21494 (two males, Malaysia); ZMB 49218
(female, Pulau Pinang, Malaysia); FMNH 171833 & 191200–201; MNHP 6897 (type); ZMB 704; ZFMK 21495, 25692,
32278, 46049 & 51162
Gonocephalus beyschlagi ZMB 48879 (male); FMNH 209727 (male) & FMNH 209728, 209729, 209731, 209733–35
(females); all Bukit Lawang, Sumatra, Indonesia; ZMB 12027 & 28619
Gonocephalus bornensis RMNH 3043/3044 (two females, types, Borneo); ZFMK 36513 (male, Poring, Sabah, Borneo,
Malaysia) / ZFMK 50527 & 50528 (one male, one female / types of G. d e n z e r i ); ZMB 9032 & 9067 (two males, Kapuas,
Central Borneo, Indonesia); FMNH 63710–713, 76269, 76270, 131548–550, 149449, 149450, 149453, 149456, 149458,
239341, 246284, 246288–290, 246293, 246294–300; ZFMK 40062; ZMA 11585 & 18933; ZRC 2888–90, 2915 & 2930
Gonocephalus chamaeleontinus ZFMK 26676 & 52823 (two males, Pulau Tioman, Malaysia); ZMB 48925 & 49220 (two
females, Pulau Tioman, Malaysia); ZMA 14053, 18896, 18897 & 18947; ZMB 706, 3007, 8463, 14396, 31276,31278,
36955 & 49219; SMF9752, 9753 & 9756; ZFMK 45140; BMNH 1933.6.20.18 & 1960.1.4.15; MNHP 1889303–0305
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Gonocphalus doriae FMNH 139589 & 188600 (two males, Nanga Tekalit, Sarawak, Borneo, Malaysia); FMNH 145831
(female, Nanga Tekalit, Sarawak, Borneo, Malaysia); FMNH 195365, 230175, 246239 & 246240; SMF 9750
Gonocephalus grandis ZFMK 19373 & 19374 (one male, one female, Malaysia); ZMB 47952 & 48830 (one female and
juveniles, one male, Pulau Tioman, Malaysia); ZMB 47950 (one semiadult male, one semiadult female, Bukit Lawang,
Sumatra, Indonesia)
Gonocephalus klossi ZMB 48800 & 48801 (one male, one female, Payakumbuh, Sumatra, Indonesia); ZMA 18932; BMNH
1946.8.1465; ZFMK 40712, 40714–18
Gonocephalus kuhlii ZMB 13060 (male, Java, Indonesia); ZMB 29014–15 (two females, Cibodas, Westjava, Indonesia); ZMB
705–3, 31273–1,2; ZMA 18899, 18926, 18927 & 18929; SMF 9754, 9755, 9757, 9758, 49721 & 49722; ZFMK 20259,
20782 & 50197; ZRC 2939; MTKD 7042
Gonocephalus lacunosus ZMB 48616 (male, holotype) ZMB 48857 (female paratype), both Berastagi, Sumatra, Indonesia;
ZMA 18930 (Berastagi, Sumatra, Indonesia); ZMA 10369 & 18931 (Sumatra)
Gonocephalus liogaster ZMB 48876 (male, Malaysia); ZMB 48877 & 49217 (two females, Malaysia); MTKD 30612 &
32572; ZMA 11584; ZMB 7111, 7112, 4995 & 15876; ZFMK 16499
Gonocephalus megalepis ZMB 49041 (male, Payahkumbuh, Sumatra, Indonesia)
Gonocephalus” Genus A mjobergi BMNH 1946.8.13.87 (female, holotype, Mt. Murud, Sarawak, Borneo, Malaysia)
Gonocephalus semperi ZMB 5377 & 54501 (syntypes, male and female, Philippines)
Gonocephalus sophiae ZMB 48648 (one female, three juveniles, Philippines)
APPENDIX 2. GenBank accession number.
16S rRNA sequences used for the present publication; where this material had been used in previous studies this is indicated.
Coryphophylax subcristatus EU502992
Gonocephalus chamaeleontinus AB070379 (Honda et al. 2002; Pyron et al. 2013)
Gonocephalus grandis AB031983 (Honda et al. 2002; Pyron et al. 2013)
Gonocephalus miotympanum (= bornensis) AB070380 (Honda et al. 2002)
Gonocephalus robinsonii AB070381 (Honda et al. 2002; Guha & Kashyap 2006; Pyron et al. 2013)
Acanthosaura armata AB266452 (Okajima & Kumazawa 2010)
Acanthosaura crucigera AB031980 (Honda et al. 2002; Pyron et al. 2013)
Aphaniotis fusca AB023771 (Honda et al. 2000; Honda et al. 2002; Pyron et al. 2013)
Bronchocela cristatella EU503024 (Pyron et al. 2013)
Calotes versicolor AB031981 (Honda et al. 2002; Pyron et al. 2013)
Draco melanopogon AB023761 (Honda et al. 2002) (AB023762 Pyron et al. 2013)
Draco volans AB023760; AB023770 (Honda et al. 2000; Honda et al. 2002; Pyron et al. 2013)
Hydrosaurus amboinensis AB475096
Hypsilurus “godeffroyi” AB031984 (Honda et al. 2002)
Japalura luei JN098482
Japalura polygonata AB031985 (Honda et al. 2002; Pyron et al. 2013)
Phoxophrys nigrilabris AB031988 (Honda et al. 2002; Pyron et al. 2013)
Physignathus cocincinus AB031990 (Honda et al. 2002, Pyron et al. 2013)
Ptyctolaemus (= Mantheyus) phuwuanensis AB023772 (Honda et al. 2000; Honda et al. 2002; Pyron et al. 2013)
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