The phylogeny of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with a description of a new species from the Mangin Range, Sagaing Region, northern Myanmar

Abstract

Phylogenetic analyses of the 49 nominal species of the gekkonid genus Hemiphyllodactylus based on the mitochondrial gene NADH dehydrogenase subunit 2 and its flanking tRNAs resulted in a strongly supported tree composed of a number of regionally localised mono-phyletic lineages consistent with previous genus-wide analyses. One such lineage from Western Indochina is composed of three previously recognised clades plus an additional species from southern Thailand. An integrative taxonomic analysis of one of these clades (clade 3 from Western Yunnan, China) recovered a new species from the Mangin Mountain Range that represents the third independent origin of Hemiphyllodactylus in Myanmar (not counting the widespread parthe-nogenetic H. typus). Hemiphyllodactylus zalonicus sp. nov. from Mt. Zalon, Sagaing Region, is the first species of Hemiphyllodactyus known from the northern part of Ayeyarwady Basin and bears a 15.0-18.9% uncorrected sequence divergence from other clade members as well as having discretely non-overlapping meristic and mensural differences. This discovery brings the total number of Hemiphyllodactylus in Myanmar to at least 11 species.

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The phylogeny of Hemiphyllodactylus Bleeker, 1860
(Squamata: Gekkonidae) with a description of
a new species from the Mangin Range, Sagaing
Region, northern Myanmar
L. Lee Grismer , May Thu Chit , Parinya Pawangkhanant , Roman A. Nazarov ,
Than Zaw & Nikolay A. Poyarkov
To cite this article: L. Lee Grismer , May Thu Chit , Parinya Pawangkhanant , Roman A. Nazarov ,
Than Zaw & Nikolay A. Poyarkov (2020) The phylogeny of Hemiphyllodactylus Bleeker, 1860
(Squamata: Gekkonidae) with a description of a new species from the Mangin Range, Sagaing
Region, northern Myanmar, Journal of Natural History, 54:29-30, 1913-1931
To link to this article: https://doi.org/10.1080/00222933.2020.1833095
Published online: 11 Feb 2021.
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The phylogeny of Hemiphyllodactylus Bleeker, 1860
(Squamata: Gekkonidae) with a description of a new species
from the Mangin Range, Sagaing Region, northern Myanmar
L. Lee Grismer
a
, May Thu Chit
b
, Parinya Pawangkhanant
c
, Roman A. Nazarov
d
,
Than Zaw
b
and Nikolay A. Poyarkov
e,f
a
Herpetology Laboratory, Department of Biology, La Sierra University, Riverside, CA, USA;
b
Department of
Zoology, University of Mandalay, Mandalay, Myanmar;
c
Rabbit in the Moon Foundation, Ratchaburi,
Thailand;
d
Zoological Museum, Moscow State University, Moscow, Russia;
e
Faculty of Biology, Department of
Vertebrate Zoology, Moscow State University, Moscow, Russia;
f
Laboratory of Tropical Ecology, Joint
Russian-Vietnamese Tropical Research and Technological Center, Hanoi, Vietnam
ABSTRACT
Phylogenetic analyses of the 49 nominal species of the gekkonid
genus Hemiphyllodactylus based on the mitochondrial gene NADH
dehydrogenase subunit 2 and its anking tRNAs resulted in a strongly
supported tree composed of a number of regionally localised mono-
phyletic lineages consistent with previous genus-wide analyses. One
such lineage from Western Indochina is composed of three previously
recognised clades plus an additional species from southern Thailand.
An integrative taxonomic analysis of one of these clades (clade 3 from
Western Yunnan, China) recovered a new species from the Mangin
Mountain Range that represents the third independent origin of
Hemiphyllodactylus in Myanmar (not counting the widespread parthe-
nogenetic H. typus). Hemiphyllodactylus zalonicus sp. nov. from Mt.
Zalon, Sagaing Region, is the rst species of Hemiphyllodactyus known
from the northern part of Ayeyarwady Basin and bears a 15.0–18.9%
uncorrected sequence divergence from other clade members as well
as having discretely non-overlapping meristic and mensural dier-
ences. This discovery brings the total number of Hemiphyllodactylus
in Myanmar to at least 11 species.
ARTICLE HISTORY
Received 9 August 2020
Accepted 1 October 2020
KEYWORDS
Integrative taxonomy;
Hemiphyllodactylus zalonicus
sp. nov.; Indochina;
phylogeny; Ayeyarwady
Basin; Burma
ZOOBANK REGISTRATION
http://www.zoobank.org/
urn:lsid:zoobank.org:pub:
A01B580C-BD55-42A7-88FE-
F2983DBE82A3
Introduction
Hemiphyllodactylus is a rapidly growing gekkonid genus due to the discovery of new
species and currently contains 49 species (Mohapatra et al. 2020; Do et al. 2020; Uetz
et al. 2020) that collectively extend from Southern India and Sri Lanka to the Western
Pacic (Grismer et al. 2013). All are small nondescript species which in Indochina are
generally restricted to circumscribed upland areas (see Grismer et al. 2013; 2014a,
2020a; Eliades et al. 2019; Sukprasert et al. 2018, and references therein).
Hemiphyllodactylus species are elusive and generally dicult to nd in their native
environments and as such, many species descriptions are based on a single or just
a few specimens (Cobos et al. 2016; Eliades et al. 2019; Grismer et al. 2013; 2014b, 2015,
CONTACT L. Lee Grismer lgrismer@lasierra.edu; Nikolay A. Poyarkov n.poyarkov@gmail.com
JOURNAL OF NATURAL HISTORY
2021, VOL. 54, NOS. 29–30, 1913–1931
https://doi.org/10.1080/00222933.2020.1833095
© 2021 Informa UK Limited, trading as Taylor & Francis Group
Published online 11 Feb 2021

2018; Grismer et al. 2018a; 2020a, 2020b; Guo et al. 2015). On the other hand, lizards may
be incredibly dense in rural situations where they are human commensals (Mohapatra
et al. 2020; Grismer et al. 2020a). At present, at least 10 species of Hemiphyllodactylus are
reliably recorded from Myanmar, though the real diversity of this genus in the region
appears to be clearly underestimated (Grismer et al. 2020a).
Indochina is considered as the major centre of Hemiphyllodactylus diversity (Grismer
et al. 2013). Our recent eldwork at Mangin Mountain Range (also known as Mhankin
Range in Burmese spelling) at Mt. Zalon (or Zalon Taung) near Ban Mauk, Sagaing Region
in northern Myanmar, resulted in the discovery of yet another new species of
Hemiphyllodactylus which represents the rst record of the genus from the northern
part of the Ayeyarwady Basin (Figure 1). Genetic data from the mitochondrial gene
NADH dehydrogenase subunit 2 (ND2) and its anking tRNAs do not place it in either
of the known distantly related Burmese clades (Grismer et al. 2020a) but align it with the
Chinese species H. changningeneis and H. longlingensis (clade 3 of Grismer et al. 2017)
from the mountains of Western Yunnan Province in southwest China. As such, this new
population represents a third Burmese lineage. However, the single specimen is vastly
dierent in morphology from H. changningeneis and H. longlingensis and therefore is
described as a new species herein.
Materials and methods
Sampling
Two specimens were collected in dipterocarp tropical forest at the foothills of Zalon
Mountain in Ban Mauk District of Sagaing Region, northern Myanmar (Figure 1).
Geographic coordinates and elevation were obtained using a Garmin GPSMAP 60CSx
and recorded in WGS 84 datum. Tissue samples were taken from one specimen (ZMMU R-
16635) and stored in 95% ethanol prior to preservation in 10% buered formalin and
Figure 1. Distribution of the species of the Western Indochina lineage of the genus Hemiphyllodactylus.
1914 L. L. GRISMER ET AL.

subsequent storage in 70% ethanol. The specimens and tissue sample were subsequently
deposited in the herpetological collections of the Zoological Museum of Moscow
University (ZMMU, Moscow, Russia) and Zoology Department of University of Mandalay
(ZDUM, Mandalay, Myanmar).
Phylogenetic analyses
We obtained 1,039 base pairs of NADH dehydrogenase subunit 2 (ND2) sequence data
from 199 specimens from GenBank (see Grismer et al. 2017, Grismer et al. 2018a, 2020a)
and a newly sequenced specimen from Sagaing, Myanmar for phylogenetic analyses. Four
outgroup species, Gehyra felmani (Taylor), G. mutilata (Wiegmann), Hemidactylus frenatus
Duméril & Bibron and Lepidodactylus lugubris Duméril & Bibron were used to root the tree
based on Heinicke et al. (2011).
For molecular phylogenetic analyses, total genomic DNA was extracted from
ethanol-preserved liver tissue using standard phenol-chloroform – proteinase
K (nal concentration 1 mg/ml) extraction procedures with consequent isopropanol
precipitation (protocols followed Hillis et al. 1996). The isolated total genomic DNA
was visualised in agarose electrophoresis in presence of ethidium bromide. The
concentration of total DNA was measured in 1 μl using NanoDrop 2000 (Thermo
Scientic) and consequently adjusted to ca. 100 ng DNA/μL.
The ND2 mitochondrial DNA gene was amplied using a double-stranded
Polymerase Chain Reaction (PCR) under the following conditions: 1.0 µl genomic
DNA (10–30 ng), 1.0 µl light-strand primer (10 µM), 1.0 µl heavy strand primer
(10 µM), 1.0 µl dinucleotide pairs (1.0 µM), 2.0 µl 5x buer (2.0 µM), 1.0 MgCl
10x buer (1.0 µM), 0.10 µl Taq polymerase (5 u/µl) and 7.4 µl H
2
O. PCR reac-
tions were executed on a Bio-Rad T100™ Thermal Cycler under the following
conditions: initial denaturation at 95°C for 2 min, followed by a second dena-
turation at 95°C for 35 s, annealing at 48°C for 35 s, followed by a cycle
extension at 72°C for 35 s, for 31 cycles. The PCR products were visualised on
a 1.0% agarose electrophoresis gel. Successful targeted PCR products were
outsourced to Evrogen® (Moscow, Russia) for PCR purication and sequencing.
Primers used for amplication and sequencing of the ND2 gene were L4437b
(5ʹ-AAGCAGTTGGGCCCATACC-3ʹ) and H5934 (5ʹ-AGRGTGCCAATGTCTTTGTGRTT
-3ʹ) (Macey et al. 1997). The obtained sequence for the specimen from Sagaing
Region was deposited in GenBank (accession number MW039150); GenBank
accession numbers for the remaining 198 specimens are listed in Grismer
et al. (2017, Grismer et al. 2018a, 2020a, 2020b).
We used maximum likelihood (ML) and Bayesian inference (BI) to estimate the phylo-
genetic relationships among the sampled geckos in our sequence alignment. An ML
phylogeny was estimated using the IQ-TREE webserver (Nguyen et al. 2015; Trinopoulos
et al. 2016) preceded by the selection of substitution models using the Bayesian
Information Criterion (BIC) in ModelFinder (Kalyaanamoorthy et al. 2017), which sup-
ported HKY+F+ G4 as the best t model of evolution for the tRNAs and TVM+F+ G4
for ND2 codon position 1, TVM+F + I+ G4 for position 2 and GTR+F + I+ G4 for position 3.
One thousand bootstrap pseudoreplicates via the ultrafast bootstrap (UFB; Hoang et al.
2017) approximation algorithm were employed and nodes having ML UFB values of 95
JOURNAL OF NATURAL HISTORY 1915

and above were considered highly supported (Minh et al. 2013). A Bayesian inference (BI)
analysis was carried out in Mr Bayes 3.2.3. (Ronquist et al. 2012) on XSEDE using the
CIPRES Science Gateway (Cyberinfrastructure for Phylogenetic Research; Miller et al. 2010)
employing GTR+I + G model of evolution to all partitions using default priors. Two
independent Markov chain Monte Carlo (MCMC) simulations were performed each with
four chains, three hot and one cold. We ran the MCMC simulation for 100 million
generations, sampled every 10,000 generations and discarded the rst 10% of each run
as burn-in. Convergence and stationarity of all parameters from both runs were checked
in Tracer v1.6 (Rambaut et al. 2014) to ensure eective sample sizes (ESS) were above 200.
Post-burn-in sampled trees from both runs were combined using the sumt function in Mr
Bayes and a 50% majority-rule consensus tree was constructed. Nodes with Bayesian
posterior probabilities (BPP) of 0.95 and above were considered highly supported
(Huelsenbeck et al. 2001; Wilcox et al. 2002). After removing taxa, not in the Indochina
clade (see designation below) to which the new species belonged, MEGA7 (Kumar et al.
2016) was used to calculate uncorrected pairwise sequence divergence among and within
the species.
Morphological analyses
For the descriptive work, colour data were taken from digital images of the
holotype in life and in preservative. For purposes of comparison, the methodol-
ogy involving the evaluation of mensural and meristic characters follows Zug
(2010) and Grismer et al. (2018a, 2020a, 2020b). Mensural data were taken with
Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting
microscope on the left side of the body where appropriate. Data recovered were
snout-vent length (SVL), taken from the tip of the snout to the vent; trunk or
axilla-groin length (AG), taken from the posterior margin of the forelimb at its
insertion point on the body to the anterior margin of the hind limb at its
insertion point on the body; head length (HL), measured from the posterior
margin of the retroarticular process of the lower jaw to the tip of the snout;
head width (HW), measured at the angle of the jaws; eye diameter (ED), the
greatest horizontal diameter of the eyeball; snout-eye length (SN), measured
from anteriormost margin of the eyeball to the tip of snout; nares-eye length
(NE), measured from the anterior margin of the eyeball to the posterior margin
of the external nares; and snout width (SW), measured between the external
nares. Meristic character states evaluated were the number of scales contacting
the nares (circumnasal scales, CN); the number of scales between the suprana-
sals (=postrostrals or intersupranasals, IS); the number of scales contacting the
medial edge of the infralabials and mental from the juncture of the second and
third infralabials on both sides (CS); the number of supralabial (SL) and infra-
labial (IL) scales counted from the largest scale immediately posterior to the
dorsal inection of the posterior portion of the upper jaw to the rostral and
mental scales, respectively; the number of longitudinal rows of ventral scales at
mid-body contained within one eyeball diameter (VS); the number of longitu-
dinal rows of dorsal scales at mid-body contained within one eyeball diameter
(DS); the number of subdigital lamellae wider than long on the rst nger (FL1)
1916 L. L. GRISMER ET AL.

and toe (TL1); lamellar formulae determined as the number of U-shaped, sub-
digital lamellae (split and single) on the digital pads of digits II–V of the hands
and feet; the total number of femoral pores (FP), precloacal pores (PP) or
femoroprecloacal pores (i.e. the contiguous rows of femoral and precloacal
pore-bearing scales [FPC]; given that the holotype is a female, the enlarged
pitted homologous scales in the proximal femoral and precloacal regions were
counted); and the number of cloacal spurs on each side of the hemipenal
swelling or immediately posterior to the vent in females. Colour pattern char-
acters evaluated were the presence or absence of dark pigmentation in the
gonadal tracts and caecum; conguration of the dark markings on the dorsum
(dorsal pattern); presence or absence of a dark pre- and/or postorbital stripe
extending to at least the neck; the presence or absence of a linear series of
white postorbital and dorsolateral spots on the trunk; and the presence or
absence of light-coloured, anteriorly projecting arms of the light-coloured post-
sacral marking.
Species delimitation
The general lineage concept (GLC: De Queiroz 2007) adopted herein proposes
that a species constitutes a population of organisms evolving independently of
other such populations owing to a lack of gene ow. By ‘independently’, it is
meant that new mutations arising in one species cannot spread readily into
another species (Barraclough et al. 2003; De Queiroz 2007). Under the GLC
implemented herein, molecular phylogenies were used to recover monophyletic
mitochondrial lineages of individual(s) (i.e. populations) in order to develop
initial species-level hypotheses – the grouping stage of Hillis (2019). Discrete
colour pattern data and morphological data were then used to search for
unique characters and patterns consistent with the previous designations of
the species-level hypotheses – the construction of boundaries representing
the hypothesis-testing step of Hillis (2019) – thus providing independent diag-
noses to complement the molecular analyses.
Results
The ML and BI analyses returned well supported, nearly identical topologies for all the
known species of the genus that were concordant with genus-wide phylogenies in
previous analyses (Grismer et al. 2013, 2017, 2018a; 2020a, 2020b). The Mt. Zalon speci-
men was recovered as a member of the typus group and embedded within a lineage of
species from Western Indochina where it is nested in clade 3 with Hemiphyllodactylus
longlingensis and H. changningensis to which it is the well-supported (BI 1.00/ML 94) sister
species of the latter (Figure 2). The Mt. Zalon specimen and H. chingningensis dier by
having an uncorrected pairwise sequence divergence of 15.0–15.3% (Table 1) and also
dier in a number of discrete (i.e. non-overlapping ranges) of meristic and mensural
characters (see comparisons). As such we hypothesise this specimen represents a new
evolutionary lineage and describe it as a new species below.
JOURNAL OF NATURAL HISTORY 1917

Figure 2. Maximum likelihood consensus tree of Hemiphyllodactylus with Bayesian posterior prob-
abilities (BPP) and Ultrafast Bootstrap support (UFB) value coding at the nodes. Clade nomenclature
follows Grismer et al. (2017, Grismer et al. 2018a, 2020a).
1918 L. L. GRISMER ET AL.

Taxonomy
Family GEKKONIDAE Gray 1825
Genus Hemiphyllodactylus Bleeker, 1860
Hemiphyllodactylus zalonicus sp. nov. (Figure 3–4; Table 2)
ZooBank registration: http://www.zoobank.org/urn:lsid:zoobank.org:act:D2B420D6-
B39E-4E2C-A8F9-8FAA3C443FB0
Holotype
Adult female (museum cat no. ZMMU R-16635; eld ID NAP-09556) collected from the
evergreen dipterocarp tropical forest on the slope of Zalon Mountain (Zalon Taung), ca.
1 km westwards from the Zalon Taung Pagoda, Zalon Taung National Forest, ca. 13 km
northwards from Ban Mauk town, Ban Mauk District, Sagaing Region, northern Myanmar
(GPS data N 24.51628°, E 095.81705° WGS; elevation 690 m a.s.l.) on 2 August 2019, at
18:00 h by P. Pawangkhanant, May Thu Chit and N.A. Poyarkov.
Paratype
Adult female (museum cat no. ZDUM-2019.12.29–038; eld ID MTZ-00038), collected from
a rock at the same locality as the holotype on 29 December 2019, at 23:30 h by May Thu Chit.
Diagnosis
Hemiphyllodactylus zalonicus sp. nov. can be distinguished from all other congeners by
possessing the unique combination of having a maximum SVL of 37.7 mm, trunk not
particularly elongate or gracile (AG/SVL ratio 0.49); eight to ten chin scales; enlarged post-
mentals; ve circumnasal scales; four intersupranasals (= postrostrals); 10 supralabials; eight
infralabials; 18 longitudinal rows of dorsal scales and nine longitudinal rows of ventral scales at
midbody contained within one eye diameter; 3333 digital formulas on hands and 3444
formulas on the feet; three subdigital lamellae on the rst nger and four subdigital lamellae
on the rst toe; perforated femoroprecloacal scales; 16 perforated precloacal scales that
extend part-way onto the proximal femoral region; one cloacal spur on each side; no plate-
like subcaudal scales; a dark postorbital stripe extending to at least base of neck; no
Table 1. Uncorrected pairwise sequence divergences between the mitochondrial ND2 lineages of
Hemiphyllodactylus in clade 3.
Species 1 2 3 4 5 6 7 8 9 10
1. H. zalonicus sp. nov. ZMMU R-16635 *
2. H. changningensis isolate YNCN06 0.150 *
3. H. changningensis isolate YNCN12 0.153 0.003 *
4. H. changningensis isolate YNCN14 0.152 0.005 0.005 *
5. H. changningensis isolate YNCN40 0.152 0.002 0.002 0.003 *
6. H. longlingensis isolate N30 0.184 0.182 0.182 0.184 0.180 *
7. H. longlingensis isolate N31 0.189 0.184 0.184 0.185 0.182 0.012 *
8. H. longlingensis isolate N32 0.187 0.182 0.182 0.184 0.180 0.010 0.002 *
9. H. longlingensis isolate N33 0.185 0.184 0.184 0.185 0.182 0.008 0.003 0.002 *
10. H. longlingensis isolate N34 0.145 0.013 0.013 0.015 0.012 0.182 0.187 0.185 0.184 *
JOURNAL OF NATURAL HISTORY 1919

dorsolateral light-coloured spots or dark dorsolateral stripe on trunk; no dark ventrolateral
stripe on trunk; dark paravertebral markings on trunk; no light-coloured postsacral marking
bearing anteriorly projecting arms; and caecum and gonads pigmented. These characters are
scored across all species in the Indochina clade, all Thai species and all other species of
Hemiphyllodactylus from Myanmar in Grismer et al. (2017, Grismer et al. 2018a, 2020a, 2020b),
Sukprasert et al. (2018), and Eliades et al. (2019).
Figure 3. Adult female holotype of Hemiphyllodactylus zalonicus sp. nov. (ZMMU R-16635) from Zalon
Mountain, Zalon Taung National Forest, Ban Mauk District, Sagaing Region, northern Myanmar, in pre-
servative. (a) Dorsal view of body. (b) Lateral view of top head. (c) Dorsal view of top of head. (d) Gular region
showing mental, postmental and chin scales arrangement. (d) Lateral view of head. (e) Precloacal and
proximal femoral regions showing arrangement of enlarged, pitted scales. Photos by Roman A. Nazarov.
1920 L. L. GRISMER ET AL.

Figure 4. Type specimens of Hemiphyllodactylus zalonicus sp. nov. from Zalon Mountain, Zalon Taung
National Forest, Ban Mauk District, Sagaing Region, northern Myanmar, in life. Holotype (ZMMU
R-16635, female): (a) General dorsolateral view. (b) Ventral view of tail. (c) Lateral view of head.
Paratype (ZDUM-2019.12.29–038, female): (d) General dorsolateral view. Photos by Parinya
Pawangkhanant (a–c) and May Thu Chit (d).
JOURNAL OF NATURAL HISTORY 1921

Table 2. Meristic, mensural (in mm) and colour pattern data from the type series of Hemiphyllodactylus zalonicus sp. nov., H. longlingensis and H. changninensis of
clade 3. Data for latter two species come from (Zhou et al. 1981) and Guo et al. (2015), respectively. Values in bold are potentially diagnostic with respect to
Hemiphyllodactylus zalonicus sp. nov. (/) = data unavailable.
Character zalonicus sp. nov. zalonicus sp. nov. longlingensis changningensis
Specimen ID ZMMU R-16635 (holotype) ZDUM-2019.12.29–038
(paratype)
Sex and age class Adult female Adult female
Chin scales (CS) 8 10 7–9 7 or 8
Postmentals distinctly enlarged Yes Yes Yes Yes
Circumnasal scales (CN) 5\5 5\5 4 or 5 3 or 4
Intersupranasals (IS) 4 3 1–3 2 or 3
Supralabial scales (SL) 10 10 9 or 10 8–11
Infralabial scales (IL) 8 9 8–10 8–10
Dorsal scales (DS) 18 17 10–14 11–15
Ventral scales (VS) 9 10 6 or 7 6–8
Lamellar formula on hand 3333 3333 3444(3)
444(5)4
33(4)3(4)
3444
Lamellar formula on foot 3444 3444 44(5)4(5)4 3444
3333
Subdigital lamellae on first finger 3 3 4 or 5 3 or 4
Subdigital lamellae on first toe 4 4 4–6 3 or 4
Precloacal and femoral pore series separate or continuous Perforated scales in continuous row Perforated scales in continuous row Continuous Continuous
Total femoroprecloacal pores 16 perf. scales 20 perf. scales 16–27 19–22
Cloacal spurs on each side 1\1 1\1 1 or 2 1 or 2
Subcaudals enlarged, plate-like No No No No
Dark postorbital stripe Yes Yes Yes Yes
Adult females yellow No No No No
Dorsolateral light-coloured spots on trunk No No No No
Dark dorsolateral stripe on trunk No No No No
Dark ventrolateral stripe on trunk No No No No
Wide vertebral area generally unicolour No No No No
Dark dorsal transverse blotches/bands No No Variable No
Dark reticulate pattern on dorsum No No Variable Yes
Dark transverse zig-zag pattern on dorsum No No variable No
Dark paravertebral markings on body Yes Yes Variable No
Postsacral marking bearing light-coloured anteriorly projecting arms No No Yes No
Caecum pigmented Yes Yes (/) (/)
Gonads pigmented Yes Yes (/) (/)
(Continued)
1922 L. L. GRISMER ET AL.

Table 2. (Continued).
Character zalonicus sp. nov. zalonicus sp. nov. longlingensis changningensis
SVL 36.4 37.7 (/) (/)
TL 25.7 34.0 (/) (/)
AG 18.1 18.9 (/) (/)
HL 8.5 8.4 (/) (/)
SN 3.4 3.5 (/) (/)
HW 5.8 5.7 (/) (/)
NE 2.7 2.4 (/) (/)
ED 2.0 2.6 (/) (/)
SW 1.2 1.2 (/) (/)
AG/SVL 0.49 0.50 0.47–0.52 0.46–0.51
HL/SVL 0.23 0.22 0.22–0.24 0.22–0.25
HW/SVL 0.15 0.15 0.17–0.19 0.17–0.18
NE/HL 0.31 0.28 0.42–0.45 0.41–0.49
ED/HL 0.23 0.30 0.22–0.25 0.21–0.25
JOURNAL OF NATURAL HISTORY 1923

Description of holotype
Adult female SVL 36.4 mm (Figure 3a); head triangular in dorsal prole (Figure 3c),
depressed, distinct from neck; lores at to slightly convex; rostrum moderate in length
(NE/HL 0.13); prefrontal region weakly concave; canthus rostralis smoothly rounded,
barely discernable; snout moderate, rounded in dorsal prole; eye large; ear opening
elliptical, small (Figure 3b); eye-ear distance greater than diameter of eye; rostral wider
than high, bordered posteriorly by large supranasals (Figure 3c); four equally sized
intersupranasals (= postnasals); external nares bordered anteriorly by rostral, dorsally by
supranasal and one internasal, posteriorly by two postnasals, ventrally by rst supralabial
(= circumnasals); 10 (R,L) rectangular supralabials tapering to below posterior margin of
eye; nine (R,L) rectangular infralabials not tapering to below posterior margin of eye;
scales of rostrum, lores, top of head, and occiput small, raised, those of rostrum largest;
dorsal superciliaries at, mostly square, subimbricate, largest anteriorly; mental triangular,
bordered laterally by rst infralabials and posteriorly by two postmentals (Figure 3d);
postmentals in contact with rst infralabial and bordered laterally by a slightly smaller
chin shield; four or ve lateral chin shields; gular scales small, subimbricate, grading
posteriorly into slightly larger, subimbricate throat and even larger pectoral scales
which grade into slightly larger, subimbricate ventrals.
Body moderate in stature, trunk not noticeably elongate (AG/SVL 0.49), dorsoven-
trally compressed; ventrolateral folds absent; dorsal scales small, granular, 18 dorsal
scales at midbody contained within one eye diameter; ventral scales at, subimbri-
cate much larger than dorsal scales, nine ventral scales contained within one eye
diameter; precloacal scales larger than abdominal scales; 16 perforated, slightly
enlarged femoroprecloacal scales in an angular series, scale at apex lacks a pit; single
enlarged tubercle (spur) on lateral margin of tail base (Figure 3e); forelimbs short,
robust in stature, covered with at, subimbricate scales dorsally and ventrally; palmar
scales slightly raised, subimbricate; all digits except digit I well developed; digit
I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and
U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula
of digits II–V 3333 (R,L); three transversely expanded lamellae on digit I; claws on
digits II–V well developed, unsheathed; distal portions of digits strongly curved,
terminal joint free, arising from central portion of lamellar pad; hind limbs short,
more robust than forelimbs, covered with slightly raised, juxtaposed scales dorsally
and by larger, at subimbricate scales anteriorly and ventrally; plantar scales slightly
raised, subimbricate; all digits except digit I well developed; digit I vestigial, clawless;
distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae
proximal to these transversely expanded; distal lamellar formula of digits II–V 3444
(R,L); four transversely expanded lamellae on digit I; claws on digits II–V well devel-
oped, unsheathed; distal portions of digits strongly curved, terminal joint free, arising
from central portion of lamellar pad; tail original, 25.7 mm in length, caudal scales
occurring in whorls; dorsal caudal scales larger than dorsal body scales, at, sub-
cycloid, subimbricate; ventrolateral caudals slightly enlarged, weakly ared anteriorly;
subcaudals at, slightly larger than dorsal caudals, not plate-like. Morphometric data
are presented in Table 2.
1924 L. L. GRISMER ET AL.

Figure 5. Habitat of Hemiphyllodactylus zalonicus sp. nov. at the type locality: Zalon Mountain, Zalon
Taung National Forest, Ban Mauk District, Sagaing Region, northern Myanmar. (a) Zalon Mountain with
Zalong Taung Pagoda. (b) Mixed semi-deciduous dipterocarp tropical forest on the Western slope of
Zalon Mountain. Photos by Parinya Pawangkhanant.
JOURNAL OF NATURAL HISTORY 1925

Colouration in life
Dorsal ground colour light-brown except for vertebral region which is brownish grey
(Figure 4a); top of head bearing dark-coloured, irregularly shaped markings; dark-
coloured, diuse stripe extends from rostral scale to forelimb insertion (Figure 4c); large,
dark-coloured, diuse, irregularly shaped, paravertebral marking on trunk; indistinct dark-
coloured markings on ventral margin of trunk becoming more distinct posteriorly; dark-
coloured, irregularly shaped markings on limbs; indistinct light-coloured postsacral band
with broken, light-coloured anteriorly projecting arms; ve indistinct, dark-coloured,
caudal bands; gular, pectoral and abdominal regions beige with dark stippling; stippling
dense in pelvic region and underside of limbs; subcaudal region reddish-orange bearing
scattered black and white scales (Figure 4b). Colouration after a year in preservative much
the same but not nearly as distinct, bright yellowish and reddish tints faded and overall
colouration turned greyish-brown (Figure 3).
Variation
Measurements and counts of the paratype female ZDUM-2019.12.29–038 are presented in
Table 2, photographs of the paratype in life are shown in Figure 4d. Morphologically, the
paratype resembles the holotype in all major characteristics. ZDUM-2019.12.29–038 has
a bright orange-red ventral surface of tail, a slightly more pronounced dark postocular
stripe which continues posteriorly after the axilla; and more irregular dark-brown blotches
on the dorsum (Figure 4d).
Distribution
Hemiphyllodactylus zalonics sp. nov. is at present known only from the type locality of
Zalon Mountain, Ban Mauk District, Sagaing Region, northern Myanmar (Figure 1). Zalon
Mountain (941 m a.s.l. at its peak) located in the center of Zalon Taung National Forest
reserve, belongs to the northern portion of the Mangin Mountain Range – a low
mountain chain located between the valleys of the Ayeyarwady (Irrawaddy) and
Chindwin rivers, occupying the northern part of Sagaing Region and the southwest
corner of Kachin Region in northern Myanmar. The actual extent of distribution of the
new species is unknown, though its occurrence in other parts of the Mangin Range is
anticipated.
Natural history
The holotype was collected at sundown (18:00 h) from a trunk of Diospyros burmanica
Kurz (ca. 1.5 m above the ground), within the primarily dipterocarp tropical semi-
deciduous forest at the foothill of Zalon Mountain (Figure 5a). The semi-deciduous forests
around Zalon Mt. are composed of broad-leaved trees such as Dipterocarpus alatus Roxb.
ex G.Don., Shorea assamica Dyer in J. D. Hooker, Dyospiros burmanica, Swintonia oribunda
Gri., Engelhardtia spicata Lesch ex Blume, Magnolia champaca (L.) Baill. ex Pierre, Schima
sp., Terminalia alata Heyne ex Roth, Quercus lamellosa Sm., Castanopsis sp., Elaeocarpus
sp., Ficus spp. and others (Figure 5b). Holotype was collected while it was crawling on the
1926 L. L. GRISMER ET AL.

bark on tree trunk, hiding in bark crevices. There are several huts in the area where the
new species was recorded; however, we have not seen other specimens on these build-
ings. The paratype was found at night (23:30 h) ca. 1 m above the ground on a large
granite rock at the same locality as the holotype. Both are gravid females.
Etymology
The specic epithet zalonicus is a Latinised toponymic adjective in nominative singular
given in reference to the type locality of Mt. Zalon (Zalon Taung) in Sagaing Region of
northern Myanmar. We suggest the following common names: Mt. Zalon Slender Gecko
(English), Zalontaung Ein Myaung Twal (Burmese).
Comparisons
The molecular analyses indicate that Hemiphyllodactylus zalonicus sp. nov. is nested within
Hemiphyllodactylus clade 3 of Yunnan, China with H. changningensis and H. longlingensis
from which it diers by an uncorrected pairwise sequence divergence of 15.0–15.3% and
14.5–18.9%, respectively (see Table 2 for comparative data). Hemiphyllodactylus zalonicus
sp. nov. is most closely related to H. changningensis but diers from it by having 17–18 as
opposed to 11–15 dorsal scales; and a narrower head (HW/SVL = 0.15 versus 0.17–0.18)
and shorter snout (NE/HL = 0.28–0.31 versus 0.41–0.49). It diers from H. longlingensis by
having more dorsal scales (17–18 versus 10–14), 16–20 perforated femoroprecloacal
scales in the female versus 19–22 femoroprecloacal pores in males; and a narrower
head (HW/SVL = 0.15 versus 0.17–0.19) and shorter snout (NE/HL 0.28–0.31 versus 0.42–-
0.45) (Table 2). These characters are scored across all species in the Indochina clade, all
Thai species and all other species of Hemiphyllodactylus from Myanmar in Grismer et al.
(2017, Grismer et al. 2018a, 2020a, 2020b), Sukprasert et al. (2018), and Eliades et al. (2019).
Discussion
A resurgence of herpetological eldwork in Myanmar has resulted in a dramatic increase
in the descriptions of new gekkonid lizards in the genera Cyrtodactylus Gray and
Hemiphyllodactylus Bleeker (Uetz et al. 2020). Many of these descriptions are not just of
single species but of entirely new clades containing multiple species, thus underscoring
the still unexplored nature of vast regions of Myanmar (Grismer et al. 2017, 2018a, 2018b,
2018c, 2018d, 2018e, 2019a, 2019b, 2020a, 2020c).
The discovery of Hemiphyllodactylus zalonicus sp. nov. brings the total number of
Hemiphyllodactylus in Myanmar to at least 11, though this is still likely an underestimate
of the diversity of this genus given its high degree of site-specic endemism and that
many upland areas remain not surveyed.
Isolated hilly areas within the Ayeyarwady Basin, including the Mangin and Kumon
Ranges, are proving to be areas of gekkonid and, more generally, herpetofaunal ende-
mism (Bauer 2002; 2003; Grismer et al. 2018d; 2019b, 2019c; Than Zaw et al. 2019;
Poyarkov et al. 2019). Hemiphyllodactylus zalonicus sp. nov. is the most recently described
species belonging to clade 3 (Figure 2) within a larger lineage from Western Indochina. It
not only represents the rst record of this genus from the Ayeyarwady Basin but repre-
sents the third independent origin of Hemiphyllodactylus in Myanmar (Figures 1, 2). The
JOURNAL OF NATURAL HISTORY 1927

south lineage (sec. Grismer et al. 2020a) is a newly described clade of species endemic to
the southern extreme of the Shan Plateau and the northern end of the Salween Basin,
Myanmar. Clade 4 (sec. Grismer et al. 2020a) is composed of six Burmese species endemic
to the Shan Plateau and another from Chiang Mai, Thailand (H. chiangmaiensis) and one
species (H. jinpingensis) scattered throughout mountains of southern and south-Western
China. Large distribution gaps between closely related species in the three Western
Indochinese clades suggest that many more species have yet to be discovered through-
out the vast upland regions of Indochina.
The area around Zalon Mountain was recently regarded as a proposed Zalong Taung
National Park, which should encompass the eastern slopes of Mangin Range north of
Banmauk town, along with hilly catchment areas of Nam Ke Hu, Chaung Gyi and Hwe
Taung Rivers between latitudes 24°27ʹN to 24°41ʹN and longitudes 95°45ʹE to 95°55ʹE. The
Zalon Taung Pagoda, located on the top of Zalon Mountain (Figure 5a), is an important site
for religious pilgrimage, and the number of local pilgrims has increased signicantly in recent
years, from 200,000 pilgrims in 2017 to over 400,000 pilgrims in 2018. In coming years more
tourists will be attracted to this area (https://elevenmyanmar.com/). The planned develop-
ment of tourist infrastructure in the area might lead to an increase of habitat loss and
modication. Further intensied survey eorts and biodiversity assessments are urgently
required for eective conservation management of yet unrealised herpetofaunal diversity of
the Zalon Mountain area.
Acknowledgements
We thank the Ministry of Natural Resources and Environmental Conservation Forest Department for
the collection and export permits. We are grateful to Mr U Thaw Moon Ko from Ban Mauk Forestry
Department for support and granting access to the Zalon Taung National Forest. We are deeply
grateful to the sta of Friends of Wildlife Association in Ban Mauk and especially to Mr Win Ko Ko
Naing Tun, Mr Wai Yan Tun and Mr Soe Htike Aung for support and help with the organization of
eldwork. We thank Mr U Ba La and Daw Aye Thar and their family for their hospitality during our
eldwork in Zalon Taung. We are grateful to Vladislav A. Gorin and Evgeniy S. Popov for help and
support during the eldwork; we thank Platon V. Yushchenko and Anna S. Dubrovskaya for
assistance during lab work. We thank Valentina F. Orlova (ZMMU) for permission to study specimens
under her care and permanent support, and to Chatmongkon Suwannapoom for support. We are
sincerely grateful to two anonymous reviewers for their kind help and useful comments, which
helped us to improve the previous version of this manuscript. May Thu Chit is grateful to Mars
Dragon Company Limited, and personally to Mr Soe Tun Zaw, for nancial support of her eldwork
in Zalon Taung National Forest within the frameworks of her PhD research project. The research was
carried out within the frameworks of Russian State projects AAAA-A16116021660077-3 and АААА-
А17-117030310017-8 (specimen storage).
Authors’ contributions
NAP and LLG designed the study. MTC, TZ, PP, RAN and NAP collected data. NAP performed
molecular analysis. RAN, MTC and NAP examined morphology. NAP and LLG supervised the
analyses. LLG and NAP wrote the manuscript. MTC, TZ, RAN and NAP revised the manuscript. All
authors read and approved the nal manuscript.
1928 L. L. GRISMER ET AL.

Disclosure statement
No potential conict of interest was reported by the authors.
Funding
This work was supported by the Russian Science Foundation [19-14-00050].
ORCID
L. Lee Grismer http://orcid.org/0000-0001-8422-3698
Parinya Pawangkhanant http://orcid.org/0000-0002-0947-5729
Nikolay A. Poyarkov http://orcid.org/0000-0002-7576-2283
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