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Discovery of Japalura chapaensis Bourret, 1937 (Reptilia: Squamata: Agamidae) from Southeast Yunnan Province, China

  • Chengdu Institute of Biology CAS

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Due to a paucity of surveys in northern Indochina and lack of international collaborations among neighboring countries, recognized distributional ranges for many amphibian and reptile species end at the political borders for some countries, despite seemingly continuous suitable habitat spanning the region. Combining both morphological and genetic data, we report the first discovery of Japalura chapaensis, a rare agamid lizard believed previously to be endemic to northern Vietnam only, along the border region of southeastern Yunnan Province, China. To facilitate future research on the genus Japalura sensu lato in Indochina, we provide detailed descriptions of additional specimens of this rare species, including the first description of coloration in life and an expanded diagnosis, and discuss the species boundary of J. chapaensis with respect to its congeners.
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Science Press Zoological Research 39(2): 105-113, 2018
Discovery of Japalura chapaensis Bourret, 1937
(Reptilia: Squamata: Agamidae) from Southeast
Yunnan Province, China
Kai Wang1,*, Ke Jiang2,3, Yu-Fan Wang4, Nikolay A. Poyarkov Jr.5,6, Jing Che2,3, Cameron D. Siler1
1 Sam Noble Oklahoma Museum of Natural History & Department of Biology, University of Oklahoma, Norman Oklahoma 73072-7029,
2 Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming Yunnan 650223, China
3 Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, Yezin Nay Pyi Taw 05282, Myanmar
4 Zhejiang Forest Resource Monitoring Center, Hangzhou Zhejiang 310020, China
5 Department of Vertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Leninskiye Gory, GSP-1, Moscow 119234,
6 Joint Russian-Vietnamese Tropical Research and Technological Center, 63 Nguyen Van Huyen Road, Nghia Do, Cau Giay, Hanoi, Vietnam
Due to a paucity of surveys in northern Indochina
and lack of international collaborations among
neighboring countries, recognized distributional
ranges for many amphibian and reptile species end
at the political borders for some countries, despite
seemingly continuous suitable habitat spanning the
region. Combining both morphological and genetic
data, we report the first discovery of Japalura
chapaensis, a rare agamid lizard believed previously
to be endemic to northern Vietnam only, along the
border region of southeastern Yunnan Province,
China. To facilitate future research on the genus
Japalura sensu lato in Indochina, we provide
detailed descriptions of additional specimens of this
rare species, including the first description of
coloration in life and an expanded diagnosis, and
discuss the species boundary of J. chapaensis with
respect to its congeners.
Keywords: Draconinae; Indochina; Morphological
Variation; New record; Range extension; Systematics;
Species with distributions spanning the political jurisdictions of
multiple countries pose special challenges to biodiversity and
conservation studies. Not only are conservation assessments
and protection of this class of evolutionary lineages more
difficult, but also, variation in the understanding of regional
biodiversity can lead to conflicting or questionable taxonomic
and biogeographic patterns for widespread species. This
situation has been particularly pronounced in peripheral island
systems, including the Philippines (Esselstyn et al., 2004;
Esselstyn & Oliveros, 2010; Siler et al., 2014), eastern
Indonesia (Taylor, 2003), and Hawaii (Gillespie et al., 2012),
among others (Gillespie & Claque, 2009). However, even for
species with broad geographic distributions on continental
landmasses, the paucity of communication across political
boundaries has resulted in species being recognized as
restricted to one side of a country’s border or the other, despite
evidence of continuous distributions spanning the boundaries of
multiple countries (Dudgeon et al., 2006; Hannah et al., 2002).
This is exemplified in regions of Southeast Asia, particularly
northern Indochina. 1
The border between southern China and northern Vietnam
has posed problems for studies of amphibian and reptile
diversity. Historically, biodiversity research on the herpetofauna
across this region has been conducted independently in China
and Vietnam, with interpretation of research often slowed by
language barriers and lack of regional collaboration (Ananjeva
Received: 14 August 2017; Accepted: 06 September 2017
Foundation items: The study is supported by The National Science
Foundation (NSF) GRFP 2017216966 and EAPSI 1714006 to K.W.,
the Russian Science Foundation (RSF grant No. 14-50-00029) to
N.A.P., Animal Branch of the Germplasm Bank of Wild Species of CAS
(Large Research Infrastructure Funding) to C.J., and NSF Integrative
Organismal Systems (IOS) 1353683 to C.D.S.
*Corresponding author, E-mail:
et al., 2007; Bain et al., 2009; Yang & Rao, 2008; Zhao et al.,
1999). However, despite the similar habitat structures across
this region, and evidence of continuous distributions for
ecologically similar species of frogs and lizards (Bain et al.,
2009; Ota, 2000), much of the herpetofauna is still thought to be
endemic to one side of the political border or the other. An
example of this is the Chapa Mountain Dragon, Japalura
chapaensis Bourret, 1937.
First described as a subspecies of Japalura swinhonis
Günther, 1864, the description of J. chapaensis was based
originally on a single female specimen (MNHN 1948.45) from
Chapa (now “Sa Pa”) in Lao Cai Province, northern Vietnam,
close to the China-Vietnam border (Bourret, 1937). Ota (1989)
later elevated J. chapaensis to full species based on
examination of morphological characters, and three years later,
Ota & Weidenhöfer (1992) described the first and only male
specimen of the species (KUZ 20097) from the type locality.
However, despite some descriptive comparisons to morphologically
similar species, low sample size (two specimens of J.
chapaensis, both with incomplete tails reported) and limited
diagnostic analysis against other congeners in the region, has
limited our understanding of the phenotypic variation and
diagnostic features of J. chapaensis (Ota, 1989; Ota &
Weidenhöfer, 1992). Currently, Japalura chapaensis is known
from only two voucher specimens only, with distribution records
limited to the eastern slopes of the Hoang Lien Son mountain
ridge in northern Vietnam (Ananjeva et al., 2007; Bourret, 1937;
Cai et al., 2015; Manthey, 2010; Van Sang et al., 2009).
However, as the Hoang Lien Son Mountains continue and
extend to the southern part of the Yunnan Province in China,
thus offering similar habitats in close geographic proximity to
the type locality of J. chapaensis in northern Vietnam, it remains
possible that J. chapaensis also occurs across the border in the
southern edge of China.
In November 2015, we collected three agamid lizards from
Lvchun, Honghe Prefecture, Southern Yunnan, China, close to
the China-Vietnam border (Figure 1). Based on newly collected
genetic and morphological datasets, we confirmed the identity
of these Chinese specimens as J. chapaensis. Here, we extend
the recognized distribution of J. chapaensis into China, provide
a description of additional specimens of this rare species, and
discuss morphological variation of J. chapaensis with relation to
mainland congeners, particularly J. yunnanensis.
Figure 1 Distribution of Japalura sensu lato in northern Indochina
Stars indicate type localities for J. yunnanensis (A: Longling, Baoshan, Yunnan Province, China) and J. chapaensis (B: Sa Pa, Lào Cai Province,
Vietnam); circle shows new locality of J. chapaensis (C: Lvchun, southern Yunnan, China).
Field sampling
A total of three specimens of J. chapaensis (an adult male, an
adult female, and a sub-adult female) were collected from
Lvchun, Honghe Prefecture, southern Yunnan Province, China
on 10 November 2015 (N22.994 1°, E102.398 9°, 1 550 m
elevation, WGS 84), and a topotypic newborn was collected
from the vicinity of Tram Don mountain pass, Hoang Lien Son
National Park, Sa Pa, Lao Cai Province, Vietnam (GPS
N22.353 1°, E103.774 9°, 1 950 m elevation, WGS 84; Figure
1). After euthanasia, tissue samples were taken from the liver
and preserved in 95% ethanol, and the voucher specimens
were fixed in 10% buffered formalin and transferred to 70%
ethanol after fieldwork. All Chinese specimens were deposited
at the Museum of Kunming Institute of Zoology, Chinese
Academy of Sciences, and the Vietnamese specimen was
deposited at the Zoological Museum of Lomonosov Moscow
State University, Moscow, Russia.
Zoological Research 39(2): 105-113, 2018
We examined all morphological characters used in previous
descriptions of J. chapaensis (Bourret, 1937; Ota, 1989; Ota &
Weidenhöfer, 1992), as well as additional characters following
Wang et al. (2016). Morphometric data were recorded by JK
using a digital caliper to the nearest 0.01 mm, except for tail
length, which was recorded using a ruler to the nearest 1 mm.
The following morphological characters were examined:
snout–vent length (SVL), tail length (TAL), trunk length (TRL),
head width (HW), snout–eye length (SEL), interorbital
distance (IOD), fore-limb length (FLL), hind limb length (HLL),
Toe IV length (T4L), Finger IV subdigital lamellae count (F4S),
supralabial count (SL), infralabial count (IL), middorsal scale
count (MD), Toe IV subdigital lamellae count (T4S), number of
scales between nasal and first supralabial (NSL), supraciliary
count (SCL), number of scale rows between the sixth supralabial
and orbit circle (SOR), enlarged, conical, occipital scale count
(COS), enlarged, conical, post-tympanic scale count (PTY), and
enlarged, conical, postrictal scale count (PRS).
Summaries of specimens examined are presented in
Appendix. For comparisons with other phenotypically similar
species, morphological data were collected from type or
topotype specimens when available. In addition to the vouchered
specimens examined, morphological data of congeners were
obtained from literature (Manthey et al., 2012; Ota &
Weidenhöfer, 1992). For maximum comparability and
consistency, color description terminology followed Köhler
(2012). Museum abbreviations followed Sabaj (2016), including:
Chengdu Institute of Biology, Chinese Academy of Sciences
(CIB); Kunming Institute of Zoology, Chinese Academy of
Sciences, Kunming, China (KIZ); Department of Zoology, Kyoto
University, Kyoto, Japan (KUZ); National Museum of Natural
History, Washington D.C., USA (NMNH); Museum of
Comparative Zoology, Harvard University, Cambridge, USA
(MCZ); California Academy of Sciences, San Francisco, USA
(CAS); National Museum of France, Paris, France (MHNP); and
Zoological Museum of Lomonosov Moscow State University,
Moscow, Russia (ZMMU).
DNA sequencing and genetic divergences
We extracted total genomic DNA from the newly collected
tissues of J. chapaensis from China (KIZ 034923 and KIZ
034921), topotypic J. chapaensis (ZMMU NAP-01911), and
topotypic J. yunnanensis (CAS 242271), using a guanidine
thiocyanate extraction protocol. A fragment of 432 bp of the
mitochondrial NADH dehydrogenase subunit 2 (ND2) gene was
targeted and amplified using the primers and protocols of
Macey et al. (2000). Previously published sequence data ND2
for congeners J. splendida and J. flaviceps were obtained from
GenBank (Accession Nos.: AF128500, AF128501). Novel
sequences were deposited in GenBank (Accession Nos.:
MG214260MG214264). Uncorrected genetic distances were
calculated using PAUP v. 4.0 (Swofford, 2002).
Genetic distance and morphological characteristics of the
southern Yunnan population
For the ND2 fragment analyzed, the two individuals from
southeastern Yunnan Province are genetically identical to each
other. The southern Yunnan individuals are 1.8% divergent from
the topotypic J. chapaensis, 5.5% from the topotypic J.
yunnanensis, 14.8% from J. splendida, and 18.2% from J.
flaviceps (Table 1).
Table 1 Uncorrected genetic distances among members of sampled Japalura sensu lato using a fragment of 432 bp ND2 gene
splendida Japalura
flaviceps Japalura
yunnanensis Japalura chapaensis
(topotype) Japalura chapaensis
(southern Yunnan)
Japalura splendida
Japalura flaviceps 0.14977
Japalura yunnanensis 0.18433 0.16359
Japalura chapaensis (topotype) 0.17972 0.15668 0.0576
Japalura chapaensis (southern Yunnan) 0.18203 0.14747 0.0553 0.01843
GenBank accession Nos. are listed in the methods.
Detailed morphometric and pholidosis characteristics are
presented in Table 2. Morphological characteristics of the
individuals from southern Yunnan resemble the diagnoses of
J. chapaensis proposed by Ota (1992) (characteristics of the
type and topotype from Ota [1992] are given in
parentheses), SL 7 or 8 (7 or 8), IL 7–9 (7), MD 35–41 (35
37), and T4S 2730 (28–30). Several morphological
characteristics measured for the newly discovered
population fall outside the currently recognized range of J.
chapaensis, including relative length of Toe IV T4L/SVL
21.31%22.35% (23.7%–26.0%), relative fore-limb length
FLL/SVL 48.14%–50.79% (vs. 50.3%54.7%), and relative
hind limb length HLL/SVL 74.66%–77.81% (vs. 80.9%
81.4%). However, all characters closely match those of Ota
(1992), with the same level of intraspecific phenotypic
variation observed for other lineages in the genus Japalura
sensu lato (Ota, 2000; Wang et al., 2017). With only two
specimens known previously for J. chapaensis, it is
expected that distinct populations of the species would
possess some degree of variation among phenotypic
Table 2 Morphological and pholidosis characteristics of Japalura chapaensis
Catalog No. KIZ 034922 KIZ 034921 KIZ034923 MHNP 1948.45 KUZ 20097
Sex M F F (subadult) F M
Type Status Holotype Topotype
SVL 67.10 68.50 50.80 59.60 58.10
TAL 168 157 109 122 (?) /
HL 21.00 22.80 17.20 19.30 19.40
HW 13.80 14.40 11.30 / /
HD 12.80 12.70 9.60 / /
SEL 8.60 8.80 6.60 8.80 8.70
IOD 12.50 10.70 9.70 11.60 10.00
FLL 32.30 33.80 25.80 30.00 31.80
HLL 50.10 53.30 38.50 48.00 47.30
T4L 15.00 14.60 10.90 14.10 15.10
TRL 30.90 29.70 23.40 27.20 28.20
TAL/SVL 249.63% 228.61% 214.37% 204.70% /
HL/SVL 31.30% 33.28% 33.86% 32.38% 33.39%
HW/HL 65.71% 63.16% 65.70% / /
SEL/HL 40.95% 38.60% 38.37% 45.60% 44.85%
FLL/SVL 48.14% 49.34% 50.79% 50.34% 54.73%
HLL/SVL 74.66% 77.81% 75.79% 80.54% 81.41%
TRL/SVL 46.05% 43.36% 46.06% 45.64% 48.54%
SL 8/8 7/7 8/7 7 8
IL 8/8 8/9 8/7 7 7
SOR 3/3 3/3 3/3 3 /
NSL 0 0 0 0 0
MD 35 41 36 35 37
F4S 23/23 25/26 23/24 22/23 24/22
T4S 27/28 28/29 30/30 27/28 30/30
PTS 2/2 2/1 2/2 / /
PTY 3/3 3/2 3/2 / /
PRS 6/7 9/7 10/8 / /
Abbreviations are listed in the methods. Data of the holotype and topotype male were obtained from literature (Bourret, 1937; Ota 1989, 1992). “/”
indicates missing data from the literature, and “” indicates an incomplete tail. Tail of the holotype was recorded as complete, with its length noted in the
original description; but it was shown to be incomplete in re-descriptions by Ota (1989, 1992).
Based on genetic and morphological data, we feel confident in
the identity of the southern Yunnan population of Japalura as J.
chapaensis; a new record for agamid lizards of China. To facilitate
future study of this rare forest agamid and closely related species,
we provide a detailed description of the newly collected individuals.
Description of specimens from Yunnan (Figure 2)
KIZ 034922, adult male; KIZ 034921, adult female; KIZ 034923,
subadult female; all collected by Yufan Wang from Lvchun,
southern Yunnan, China.
Description of southern Yunnan population.—Body size
moderate, SVL 67.1 mm in adult male, 68.5 mm in adult female.
Tail slender, long, TAL/SVL 249.63% in adult male, 228.61% in
adult female, 214.37% in subadult female. Relative head length
moderate, HL/SVL 31.30% in male, 33.28% in female, 33.86%
in subadult female; HW/HL 65.71% in male, 63.16% in female,
65.70% in subadult female. Snout blunt, rounded; rostral
rectangular, three times longer than height; nasal large,
somewhat oval in shape, separated from rostral by single scale,
in contact with supralabials; SL 7 or 8, elongated, weakly keeled;
three suborbital scale rows from inferior orbit circle to sixth
Zoological Research 39(2): 105-113, 2018
Figure 2 Photographs of (1) dorsal body view, (2) ventral body view, and (3) lateral head close-ups of male (A13; KIZ 034922) and female
(B13; KIZ 034921) Japalura chapaensis in life (Photographs by Ke Jiang)
supralabial, middle row much enlarged; supraciliaries 6 or 7,
imbricate, overlapping one-fifth to one-third of posterior one;
numerous large, keeled scales forming single ridge from
postorbital to supra-anterior tympanum on each side;
tympanum concealed, covered by fine scales; two or three
large conical scales posterior to tympanum. Mental pentagonal.
IL 79, smooth; numerous large, conical postrictal scales
present. Dorsal head scales heterogeneous in size and shape,
strongly keeled; three large, hexagonal, conical scales arranged
in triangular position on dorsal snout; interparietal scale
elongate, hexagonal, parietal eye present; two pairs of distinct,
conical scales present on raised areas posterior to interparietal
on each side, forming w-shape ridge anterior to beginning of
nuchal crest; single, large, conical, postorbital scale posterior to
last supraciliaries; keeled, sub-pyramidal scales on occipital
region 2 or 3, with one distinctively tall, enlarged. Ventral head
scales keeled, more greatly keeled posteriorly, heterogeneous in
size except for a few irregularly scattered large scales, particularly
on lateral surfaces; gular pouch present, distinct in life;
longitudinal gular fold present in life; transverse gular fold absent.
Dorsal body scales strongly keeled, heterogeneous in size
and shape; scales of axillary region smaller, circular shaped;
shoulder fold present, short, weakly defined; enlarged scales
scattered on dorsal body, moderately raised, each bearing a
single distinct keel; some enlarged scales arranged in
paravertebral row close to vertebral crest; mid-dorsal crest
scales 3541, larger than neighboring scales; nuchal crest
scales 58, greatly enlarged, triangular shaped; dorsal crests
relatively lower, serrated. Ventral body scales keeled, largely
homogeneous in size; enlarged scales scattered ventrolaterally.
Limb scales keeled, largely homogeneous in size on fore-limbs,
more heterogeneous in size on hind limbs; enlarged, keeled
scales scattered irregularly on posterior hind surfaces and crus.
Tail slender, scales strongly keeled.
Coloration in life (Figure 2)
Color codes follow Köhler (2012). Both males and females of J.
chapaensis are sexually dichromatic. For males, the background
color of head varies from chamois (code 84) to olive sulphur
yellow (code 90). Four dark brownish olive (code 127) to dusky
brown (code 285) transverse bands span the dorsal surface of
the head, two of which are located on the snout, one between
the eyes and one posterior to the eyes. The background
coloration of the lateral surfaces of the head are light buff (code
2) to light flesh color (code 250). Nine dusky brown (code 285)
stripes radiate out around each eye. The two posterior-directing
radial stripes are the broadest, with the superior one extending
from the posterior corner of the eye to the anterior tympanic
region, and the inferior one extending to the corner of the
mouth. The subocular radial stripes do not extend outside of the
circular orbit. Three to four dusky brown (code 285) blotches
are scattered on both the supralabials and infralabials. The oral
cavity and tongue are dark spectrum yellow (code 79).
The background color of the neck and axillary region is light
flesh color (code 250), which transitions to olive sulphur yellow
(code 90), yellow green (code 103), and eventually citrine (code
119) posteriorly. Most regions of the dorsal and lateral surfaces
of the body possess dusky brown (code 285) reticulated
patterns around the axillary region and blotches along the
dorsal midline. An irregularly shaped dorsolateral stripe is
present on each side of the body, consisting of three Pratt’s
rufous (code 72) colored, elongated blotches, running
posterolaterally away from the vertebral midline. The coloration
of these stripes can change to chamois (code 84) depending on
the condition or mood of the lizards. Five small, sulphur yellow
(code 80) blotches are evenly scattered along the dorsal
midline from the shoulder to the pelvis. The dorsal surfaces of
the limbs are olive sulphur yellow (code 90), with numerous
dusky brown (code 285) bands spanning the upper surfaces of
the limbs to the digits. A pale buff (code 1) stripe is present on
the posterior surfaces of the upper hind limbs. The tail is pale
buff (code 1), with six citrine (code 119) bands spanning about
one-third of its total length. The bands of the tail extend to the
ventral surface, forming complete rings around the tail. The
remaining posterior surfaces of the tail are uniform ground
cinnamon (code 270) in color.
The ventral surface of the head is pale buff (code 1), with
some dusky brown (code 285) speckles scattered randomly. A
distinct dark spectrum yellow (code 78) gular spot is present in
the center of the gular region. The background color of the
ventral surface of the body and limbs is pale horn (code 11)
anteriorly, with the coloration changing to light pale pinkish buff
(code 3) posteriorly, including the upper ventral surfaces of the
limbs and the ventral surfaces of the tail. Grayish horn (code
268) speckles are scattered across the ventral surfaces of the
body and limbs.
Female body coloration differs from male coloration
(described above) as follows. The dorsal surfaces of the head
and neck are mostly uniform clay (code 18) in coloration. The
lateral surfaces of the head are yellow-green (code 103), with
two to four clay (code 18) colored radial stripes extending from
around the eyes. Five rectangular clay (code 103) blotches are
scattered evenly along the dorsal surface of the body at the
midbody, with all blotches loosely connected along the dorsal
midline. The background color of the lateral body surfaces,
dorsal limb surfaces, and dorsal tail surface are pistachio (code
102). A pale buff (code 1) stripe is present posteriorly on each
upper hind limb. Clay (code 18) bands are present on the first
one-third of the tail; however, the bands extend to the lateral
surfaces of the tail only, and do not form complete rings. The
posterior two-thirds of the tail is uniform clay (code 18).
The ventral surface of the head is uniform light pistachio
(code 101) with no speckles. This coloration transitions
gradually to dark spectrum yellow (code 79) toward the center
of the gular region, forming a gular spot. The ventral surfaces of
the body, limbs, and tail are pale green (code 99).
Ecology and distribution
Japalura chapaensis is arboreal, inhabiting low shrubs in
tropical forest, particularly near streams and along forest edges
(Figure 3). In Lvchun, individuals are usually found resting on
fern leaves at night. Currently, the species is known from
Lvchun County of Honghe Prefecture, Yunnan Province, China,
and from Cao Bang, Hai Duong, and Lao Cai Provinces of
Vietnam (Ananjeva et al., 2007). In addition, morphologically
similar individuals have been photographed in Daweishan
Natural Reserve of Honghe Prefecture, 130 km east from
Lvchun County, which suggests that the species has a wider
geographic distribution along the China-Vietnam border (Jian
Wang and Shuo Qi, personal communications).
Diagnoses from congeners
Previous studies have compared J. chapaensis with five island-
congeners (J. brevipes, J. makii, J. mitsukurii [synonym of J.
swinhonis], J. polygonata xanthostoma, and J. swinhonis), and
two mainland-congeners (J. splendida and J. yunnanensis)
(Ota, 1989; Ota & Weidenhöfer, 1992). Based on their results,
previous authors concluded that J. chapaensis is morphologically
most similar to J. brevipes, J mitsukurii, J. swinhonis, and J.
yunnanensis, but can be distinguished from these lineages by
its relative head length, relative snout-eye length, Toe IV length,
and Toe IV subdigital lamellae and middorsal crest scale counts
(Ota, 1989; Ota & Weidenhöfer, 1992).
With the addition of the new collections from southern
Yunnan and a more comprehensive examination of congeners,
we found that J. chapaensis can be readily differentiated from
all five Taiwanese species by having much larger nuchal crest
scales and a distinct head shape with a w-shaped lump on the
occipital region anterior to the nuchal crest. In addition, we
found J. chapaensis to be morphologically most similar to J.
yunnanensis, with both species lacking a transverse gular fold,
and possessing concealed tympanum, large nuchal crests,
irregularly distributed and enlarged ventrolateral scales, dark
spectrum yellow gular spots in males (sometimes in females),
and jagged dorsolateral stripes that are oriented in a
posterolateral direction away from the dorsal midline. However,
although phenotypically similar, Japalura chapaensis differs
from J. yunnanensis by having a tendency toward fewer mid-
dorsal crest scales (MD 3541 vs. 3946), and relatively shorter
tails (TAL/SVL 249.63% in male, 204.70%228.61% in females
vs. 255.19%288.81% in males, 237.45%259.67% in females)
(Table 2).
Zoological Research 39(2): 105-113, 2018
Figure 3 Habitat of Japalura chapaensis in Lvchun, southeastern Yunnan Province, China (Photograph by Yu-Fan Wang)
For the remaining mainland congeners from non-Himalayan
regions, J. chapaensis differs from J. fasciata by having a
longer tail (TAL >204% SVL vs. <180%) and longer hind limbs
(HLL >74% SVL vs. <69%), and by the absence of a transverse
hourglass-shaped marking on the dorsum (vs. present); from J.
batangensis, J. brevicauda, J. dymondi, J. flaviceps, J. grahami,
J. iadina, J. laeviventris, J. splendida, J. vela, J. yulongensis,
and J. zhaoermii by having significantly enlarged nuchal crest
scales (vs. not significantly differentiated from dorsal crest
scales), bright yellow oral cavity (vs. flash color or blackish blue),
and by the absence of a transverse gular fold (vs. present);
from J. varcoae by having a concealed tympanum (vs. exposed)
and enlarged, differentiated nuchal crest scales (vs. not
significantly differentiated); and from J. micangshanensis by
having significantly enlarged nuchal crest scales (vs. not
significantly differentiated from dorsal crest scales) and by the
presence of gular spots (vs. absent).
In conclusion, Japalura chapaensis can be diagnosed from its
congeners on the basis of the following set of characters: (1)
body size moderate SVL 58.1068.50 mm; (2) tail relatively
long TAL/SVL 249.63% (male), 204.70%228.61% (females);
(3) limb length moderate, FLL/SVL 48.14%54.73%, HLL/SVL
74.66%81.41%; (4) short, w-shaped ridge present on occipital
head anterior to nuchal crest; (5) tympanum concealed; (6)
supraciliaries overlapping less than one-third of posterior one;
(7) NSL absent (8) lateral gular fold and gular pouch present,
well developed; (9) transverse gular fold absent; (10) nuchal
crest scales large, crest height/HL 10.48%–14.04%; (11) MD
3541; (12) T4S 2830; (13) ventral scales heterogeneous in
size with enlarged scales scattered ventrolaterally; (14) ground
body coloration camouflaged with yellow-green, clay, and dusky
brown coloration in life; (15) dorsolateral stripes present in
males, jagged, sometimes discontinuous, posterolaterally away
from the dorsal midline; (16) gular spot present, distinct in
males, somewhat faded in females, dark spectrum yellow in
both sexes; and (17) oral cavity and tongue dark spectrum
yellow in life.
Despite our discovery of a new population of J. chapaensis in
southern China, the small number of individuals seen in the wild
continues to limit our understanding of this rare Japalura
species in northern Indochina. Future surveys and greater
international collaborations throughout this region are needed to
clarify the morphological variation, distribution patterns, and
population-level genetic diversity of J. chapaensis.
The authors declare that they have no competing interests.
Y.W. and N.A.P. collected the specimens. K.J. and K.W. measured
morphological data. K.W. and C.D.S. wrote the manuscript with other authors’
inputs, and C.J. revised the manuscript and gave critical feedbacks. All
authors read and approved the final manuscript.
We would like to thank Mr. Jian Xu and Shuo Qi (Shengyang Institute of
Herpetology) for providing additional locality information; Mr. Anton
Shchinov, Dr. Andrei N. Kuznetsov, and Alexandra Elbakyan for
assistance and support; and Mr. Wynn Addison (NMNH), Professor Yue-
Zhao Wang (CIB), Professor Yue-Ying Chen (CIB), Mr. Ke Lu (CIB), and
Mr. Gui-Wu He (KIZ) for kindly allowing us to examine specimens under
their care.
Ananjeva NB, Orlov NL, Truong NQ. 2007. Agamid lizards (Agamidae,
Acrodonta, Sauria, Reptilia) of Vietnam. Zoosystematics and Evolution,
83(S1): 1321.
Bain RH, Stuart BL, Nguyen TQ, Che J, Rao DQ. 2009. A new Odorrana
(Amphibia: Ranidae) from Vietnam and China. Copeia, 2009(2): 348362.
Bourret R. 1937. Notes herpétologiques sur l’indochine française. XV.
Lézards et serpents reçu au laboratoire des Sciences Naturelles de
l’Université au cours de l’année 1937. Descriptions de deux espèces et de
deux variétés nouvelles. Bulletin Générale de l’Instruction Publique 5.
Gouvernement Généneral de l’Indochine, 5782.
Cai B, Wang YZ, Chen YY, Li JT. 2015. A revised taxonomy for Chinese
reptiles. Biodiversity Science, 23(3): 365382. (in Chinese)
Dudgeon D, Arthington AH, Gessner MO, Kawabata ZI, Knowler DJ,
Lévêque C, Naiman RJ, Prieur-Richard AH, Soto D, Stiassny MLJ, Sullivan
CA. 2006. Freshwater biodiversity: importance, threats, status and
conservation challenges. Biological Reviews, 81(2): 163182.
Esselstyn JA, Oliveros CH. 2010. Colonization of the Philippines from
Taiwan: a multi-locus test of the biogeographic and phylogenetic
relationships of isolated populations of shrews. Journal of Biogeography,
37(8): 15041514.
Esselstyn JA, Widmann P, Heaney LR. 2004. The mammals of Palawan
Island, Philippines. Proceedings of the Biological Society of Washington,
117(3): 271302.
Gillespie RG, Baldwin BG, Waters JM, et al. 2012. Long-distance dispersal:
a framework for hypothesis testing. Trends in Ecology and Evolution, 27(1):
Gillespie RG, Claque DA. 2009. Encyclopedia of Islands. Berkeley, CA:
University of California Press, 11111.
Hannah L, Midgley GF, Lovejoy T, Bond WJ, Bush M, Lovett JC, Scott D,
Woodward FI. 2002. Conservation of biodiversity in a changing climate.
Conservation Biology, 16(1): 264268.
Köhler G. 2012. Color Catalogue for Field Biologists. Offenbach: Herpeton, 1
Macey JR, Schulte II JA, Larson A, Ananjeva NB, Wang YZ, Pethiyagoda R,
Rastegar-Pouyani N, Papenfuss TJ. 2000. Evaluating trans-Tethys
migration: an example using acrodont lizard phylogenetics. Systematic
Biology, 49(2): 233256.
Manthey U. 2010. TERRALOG: Agamid Lizards of Southern Asia
Draconinae 2, Leiolepidinae. Frankfurt am Main, Germany: Chimaira, 1
Manthey U, Denzer W, Hou M, Wang XH. 2012. Discovered in historical
collections: two new Japalura species (Squamata: Sauria: Agamidae) from
Yulong snow mountains, Lijiang prefecture, Yunnan, China. Zootaxa,
(3200): 2748.
Ota H. 1989. The status of an agamid lizard, Japalura swinhonis
chapaensis Bourret, 1937, from Vietnam. Journal of Herpetology, 23(4):
Ota H, Weidenhöfer T. 1992. The first male specimen of the poorly known
agamid lizard Japalura chapaensis Bourret, 1937 (Reptilia: Sauria), from
northern Vietnam, with notes on its taxonomic status. Raffles Bulletin of
Zoology, 40(2): 193199.
Ota H. 2000. Japalura szechwanensis, a junior synonym of J. fasciata.
Journal of Herpetology, 34(4): 611614.
Sabaj MH. 2016. Standard symbolic codes for institutional resource
collections in herpetology & ichthyology. American Society of Ichthyologists
and Herpetologists, USA. (20160816)
Siler CD, Oaks JR, Cobb K, Ota H, Brown RM. 2014. Critically endangered
island endemic or peripheral population of a widespread species?
Conservation genetics of Kikuchi's gecko and the global challenge of
protecting peripheral oceanic island endemic vertebrates. Diversity and
Distributions, 20(7): 756772.
Swofford DL. 2002. PAUP*: phylogenetic analysis using parsimony, version
4.0 b10. Sunderland: Sinauer Associates.
Taylor JG. 2003. Indonesia: Peoples and Histories. New Haven, London:
Yale University Press, 1–378.
Van Sang N, Cuc HT, Truongis NQ. 2009. Herpetofauna of Vietnam.
Frankfurt am Main: Edition Chimaira, 1–768.
Wang K, Jiang K, Zhou DH, Yan F, Siler CD, Che J. 2016. Two new species
of Japalura (Squamata: Agamidae) from the Hengduan Mountain range,
China. Zoological Research, 37(1): 4156.
Wang K, Ren JL, Jiang K, Yuan ZY, Che J, Siler CD. 2017. Rediscovery of
the enigmatic mountain dragon, Japalura yulongensis (Reptilia: Sauria:
Agamidae), with notes on its natural history and conservation. Zootaxa,
4318(2): 351363.
Yang DT, Rao DQ. 2008. Amphibia and Reptilia of Yunnan. Kunming:
Yunnan Publishing Group Corporation, 1411. (in Chinese)
Zhao EM, Zhao KT, Zhou KY, et al. 1999. Fauna Sinica, Reptilia, Vol. 2:
Squamata, Lacertilia. Beijing: Science Press, 1394. (in Chinese)
Specimens examined
Japalura batangensis (n=16): CIB 19021908, 2227, 2233, 2243, KIZ 84011,
801081, Batang, Sichuan, China; KIZ 09404, 019311, 019312, KIZ 019314,
Mangkang, Tibet, China.
Japalura dymondi (n=7): CIB 1869, 87234, Panzhihua, Sichuan, China; KIZ
95I1001, 1002, 1016, 1018, 1022, Dayao, Yunnan, China.
Japalura fasciata (n=3): CIB 2620, 2621, 2622, Pen Hsien, Sichuan
Province, China.
Japalura flaviceps (n=13): CIB 2234, 2332, 2333, 2341, 2354, 2355, 2549,
2554, 2556, 2561, 2567; KIZ 05181, 05182; Luding, Sichuan, China.
Japalura grahami (n=1): USMN 65500 (holotype), Yibin, Sichuan, China.
Japalura makii (n=2): MCZ R-172743 (paratype), R-181443, Taiwan, China.
Japalura micangshanensis (n=9): CIB 86348, 86351, Xianyang, Shaanxi,
China; CIB 86356, 86357, 86360, 86361, Luonan, Shaanxi, China; CIB 2572,
2578, 2582, Wenxian, Gansu, China.
Japalura polygonata (n=3): CAS 21215, 21243, 21244, Kagoshima
Prefecture, Japan.
Zoological Research 39(2): 105-113, 2018
Japalura splendida (n=6): USNM 35522 (holotype), Yichang, Hubei, China;
CIB 2588, 2591, 2596, 72468, 72469, Chongqing, China.
Japalura swinhonis (n=4): CAS 18085, 18089, 18098, 18099, Taiwan, China.
Japalura varcoae (n=3): CIB 2650, 2651, KIZ 85II0006, Kunming, Yunnan,
Japalura vela (n=11): KIZ 013801 (holotype), KIZ 013802, 013813, 013800,
013805013811 (paratopotypes), Jerkalo, Tibet, China.
Japalura yunnanensis (n=8): CIB 2684, 2686, 2687, 2689, KIZ 82081,
Longling, Yunnan, China; KIZ 74II0240, 0248, 79I469, Tengchong, Yunnan,
Japalura zhaoermii (n=14): CIB 2690 (holotype), 86432, 86435, 85721,
85722, 86433, 86434, 86436, Wenchuan, Sichuan, China; CIB 2232, 2244,
2240, KIZ 84032, 85030, Lixian, Sichuan, China.
... We compared morphological characters of the new species with other members of the genus relying on original species descriptions (Hallowell 1861;Günther 1864;Anderson 1878;Boulenger 1906Boulenger , 1918Barbour and Dunn 1919;Stejneger 1924;Mertens 1926;Smith 1935;Gressitt 1936;Bourret 1937;Song 1987;Ota 1989;Ota et al. 1998;Li et al. 2001;Gao and Hou 2002;Manthey et al. 2012;Wang et al. 2015Wang et al. , 2016Wang et al. , 2017Wang et al. , 2019bWang et al. , d, 2021aWang et al. , b, 2022Ananjeva et al. 2017;Rao et al. 2017;Liu et al. 2020) and the additional data from Wu et al. (2005), Manthey (2008) and Wang et al. (2017Wang et al. ( , 2018Wang et al. ( , 2019bWang et al. ( , c, 2021a. ...
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Three new species of Diploderma are described from the Hengduan Mountain Region in south-western China, based on morphological and genetic data. The first new species from Yulong County, Yunnan Province is morphologically most similar and phylogenetically closely related to D. brevicauda , but it can be diagnosed from the latter by having a relatively longer tail; the second new species from Xiangcheng County, Sichuan Province is phylogenetically closely related to D. bowoense , but it can be diagnosed from the latter by the absence of a distinct gular spot; and the third new species from Yongsheng County, Yunnan Province is phylogenetically closely related to D. yulongense , but it can be diagnosed from the latter by having different colourations of the ventral and ventrolateral surfaces of the body. Taxonomy and diversity survey are the basis of species conservation, our discoveries contributing to better conservation of the species of this genus.
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Although the genus Japalura s.l. has long been recognized as paraphyletic based on limited genetic sampling, its problematic taxonomy has not been revised, and phylogenetic relationships among the majority of congeners remain unknown. Here we utilize a densely sampled dataset of both multilocus genetic and morphological data to provide the first phylogenetic inference of relationships among Japalura s.l.species. Our results show that Japalura s.l. is paraphyletic, consisting of four major clades that are scattered across the phylogeny of the subfamily Draconinae: the first clade from the western, central and middle-eastern Trans-Himalayas, the second clade from the far eastern Trans-Himalayas, the third clade from East Asia and the last clade from Indochina. To address this widespread paraphyly of the genus and to stabilize the taxonomy within the family Draconinae, we revise the current taxonomy and split Japalura s.l. into four genera. By doing so, we recognize two existing generic names, Japalura sensu stricto and Pseudocalotes, resurrect one name available in the literature, Diploderma, and describe one new genus, Cristidorsa gen. nov. We discuss phylogenetic relationships and taxonomy within Japalura s.l. and present a diagnostic key to all recognized genera of the subfamily Draconinae.
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The mammal fauna of Palawan Island, Philippines is here doc- umented to include 58 native species plus four non-native species, with native species in the families Soricidae (2 species), Tupaiidae (1), Pteropodidae (6), Emballonuridae (2), Megadermatidae (1), Rhinolophidae (8), Vespertilionidae (15), Molossidae (2), Cercopithecidae (1), Manidae (1), Sciuridae (4), Muridae (6), Hystricidae (1), Felidae (1), Mustelidae (2), Herpestidae (1), Viverridae (3), and Suidae (1). Eight of these species, all microchiropteran bats, are here reported from Palawan Island for the first time (Rhinolophus arcuatus, R. ma- crotis, Miniopterus australis, M. schreibersi, and M. tristis), and three (Rhin- olophus cf. borneensis, R. creaghi, and Murina cf. tubinaris) are also the first reports from the Philippine Islands. One species previously reported from Pa- lawan (Hipposideros bicolor )i sremoved from the list of species based on re- identificaiton as H. ater, and one subspecies (Rhinolophus anderseni aequalis Allen 1922) is placed as a junior synonym of R. acuminatus. Thirteen species (22% of the total, and 54% of the 24 native non-flying species) are endemic to the Palawan faunal region; 12 of these are non-flying species most closely related to species on the Sunda Shelf of Southeast Asia, and only one, the only bat among them (Acerodon leucotis), is most closely related to a species en- demic to the oceanic portion of the Philippines. Of the 28 insectivorous bats, 18 species are somewhat to highly widespread in Indo-Australia, 2 are shared only with the Sunda Shelf and Indochina, 1 with the Sunda Shelf alone, 3 occur on the Sunda Shelf and the oceanic Philippines, 1 occurs in Palawan, Sulawesi, and the oceanic Philippines, 2 occur only on Palawan and in the oceanic Philippines, and 1 occurs on Borneo, Sulawesi, and throughout the Philippines. Though the insectivorous bats tend to be widely distributed, these data, particularly the distributions of the non-volant species, strongly reinforce the perception of Palawan Island (and associated smaller islands) as a biogeo- graphic unit of the Sunda Shelf, with only limited similarity to other portions of the Philippine Islands. The Philippine archipelago is remarkable for the large number of indigenous land mammal species (ca. 175), and especially for the number of endemic species (ca. 112). Given its relatively small land area, the Philippines has perhaps the greatest concentration of endemic mammals in the
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Being a hotspot of plant and animal diversity and endemism, Vietnam is important for the conservation of biodiversity on a global scale. It is one of the most threatened areas in tropical Asia in terms of destruction of tropical rain forest as an environment with a unique Vietnamese fauna. Previous summaries on Vietnamese agamids, their taxonomy and distribution were published in the works of Bourret 1943, Nguyen & Ho 1996, Bobrov 1995, Ziegler 2002 and Ziegler et al. 2006. New, intensive study of the fauna of Vietnam as well as new opportunities offered by molecular methods make clear that taxonomic diversity in Southeast Asia is underestimated. The aim of this paper is to show and review the taxonomic diversity of different evolutionary lineages of agamids and their geographical distribution. This is based on new results of the authors and most recent evaluation of literature sources. (© 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)
Indonesia is the fourth largest country in the world. It comprises more than 17,000 islands inhabited by 230 million people who speak over 300 different languages. Now the world's largest Muslim nation, Indonesia remains extraordinarily heterogeneous due to waves of immigration - Buddhist, Hindu, Arab, and European - that have defined the region's history." "Fifty years after the collapse of Dutch colonial rule, Indonesia is a nation in the midst of dramatic upheaval. In this broad survey, Jean Gelman Taylor explores the connections between the nation's many communities, and the differences that propel contemporary breakaway movements." "Drawing on a broad range of sources, including art, archaeology, and literature, Taylor provides a historical overview from the prehistoric period to the present day. The text is enlivened by capsule histories on topics ranging from pepper to Maharajas to smallpox." "This book - the first new history of Indonesia written in more than twenty years - will be essential reading for anyone interested in the history of Southeast Asia and the future stability of the region.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
AimTo highlight the significant conservation challenge of evaluating peripheral endemic vertebrates in island archipelago systems and to assess empirically the complexities of approaches to conservation genetic studies across political and biogeographic boundaries. To demonstrate the poignant need for international collaboration and coordination when species delimitation problems with high conservation concern involve island endemics with biogeographically peripheral ranges. LocationSoutheast Asia, Lanyu Island, Taiwan, and the Philippines. Methods Genetic samples were collected and sequenced for one mitochondrial gene and five nuclear loci for species of the Gekko mindorensis-G. kikuchii species complex in Southeast Asia. We used maximum likelihood and Bayesian phylogenetic methods and coalescent-based species delimitation analyses to estimate phylogeographic relationships, construct multilocus haplotype networks and test putative species boundaries. ResultsPhylogenetic and population genetic analyses suggest that Kikuchi's Gecko may represent a peripheral population of a widespread species distributed from the northern Philippines to Taiwan. However, we identify a discrepancy between inferences of species boundaries resulting from methods based on allele frequencies versus coalescent-based methods that incorporate evolutionary history. Coalescent-based analyses suggest that G. kikuchii may be a distinct evolutionary lineage. Our study underscores the need for coalescent-based methods in conjunction with population genetic approaches for conservation genetic assessments of widespread species. Main conclusionsThis study joins a few recent works suggesting that Philippine-derived anomalies in the fauna of Lanyu (and possibly greater Taiwan) are worthy of careful reconsideration. Determining whether each is the result of recent human-mediated introduction or (possibly more ancient) natural dispersal should be the goal of future studies on this seldom-conceived biogeographic relationship. Isolated species endemic to islands on the outer periphery of biogeographic and political regions represent particular conservation challenges. This is especially true if a species occurs on an isolated island that is allied biogeographically with one nation, but politically administered by another.