Two new species of Japalura (Squamata: Agamidae) from the Hengduan Mountain Range, China

Abstract

Until recently, the agamid species, Japalura flaviceps, was recognized to have the widest geographic distribution among members of the genus occurring in China, from eastern Tibet to Shaanxi Province. However, recent studies restricted the distribution of J. flaviceps to the Dadu River valley only in northwestern Sichuan Province, suggesting that records of J. flaviceps outside the Dadu River valley likely represent undescribed diversity. During two herpetofaunal surveys in 2013 and 2015, eight and 12 specimens of lizards of the genus Japalura were collected from the upper Nujiang (=Salween) Valley in eastern Tibet, China, and upper Lancang (=Mekong) Valley in northwestern Yunnan, China, respectively. These specimens display a unique suite of diagnostic morphological characters. Our robust comparisons of phenotype reveal that these populations can be distinguished readily from J. flaviceps and all other recognized congeners. Here-in, we describe the two Japalura lineages as new species, Japalura laeviventris sp. nov. and Japalura iadina sp. nov.. In addition, we provide updated conservation assessments for the new species as well as imperiled congeners according to the IUCN criteria for classification, discuss the importance of color patterns in the diagnosis and description of species in the genus Japalura, and discuss directions for future taxonomic studies of the group.

Full text

ZOOLOGICAL RESEARCH
Science Press Zoological Research 37(1): 41-56, 2016 41
Two new species of Japalura (Squamata: Agamidae)
from the Hengduan Mountain Range, China
Kai WANG1,2,*, Ke JIANG1, Da-Hu ZOU3,1, Fang YAN1, Cameron D. SILER2, Jing CHE1,*
1 Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming Yunnan 650223, China
2 Sam Noble Oklahoma Museum of Natural History and Department of Biology, University of Oklahoma, Norman OK 73072-7029,
U.S.A.
3 Tibet University, Lhasa Tibet 850000, China
ABSTRACT
Until recently, the agamid species, Japalura flaviceps,
was recognized to have the widest geographic dis-
tribution among members of the genus occurring in
China, from eastern Tibet to Shaanxi Province.
However, recent studies restricted the distribution of
J. flaviceps to the Dadu River valley only in north-
western Sichuan Province, suggesting that records
of J. flaviceps outside the Dadu River valley likely
represent undescribed diversity. During two herpeto-
faunal surveys in 2013 and 2015, eight and 12
specimens of lizards of the genus Japalura were
collected from the upper Nujiang (=Salween) Valley
in eastern Tibet, China, and upper Lancang
(=Mekong) Valley in northwestern Yunnan, China,
respectively. These specimens display a unique
suite of diagnostic morphological characters. Our
robust comparisons of phenotype reveal that these
populations can be distinguished readily from J.
flaviceps and all other recognized congeners. Herein,
we describe the two Japalura lineages as new spe-
cies, Japalura laeviventris sp. nov. and Japalura
iadina sp. nov.. In addition, we provide updated con-
servation assessments for the new species as well
as imperiled congeners according to the IUCN crite-
ria for classification, discuss the importance of color
patterns in the diagnosis and description of species
in the genus Japalura, and discuss directions for
future taxonomic studies of the group.
Keywords: Coloration; Conservation; Hengduan Moun-
tains; Hydropower development; Japalura flaviceps
INTRODUCTION
The family Agamidae, a radiation of more than 300 species
of iguanian lizards, is one of the most taxonomically diverse
lizard groups in Eurasia, with members of the family pos-
sessing a wide range of morphologies and life history traits
(Manthey, 2010; Moody, 1980; Townsend et al., 2011). Due
to this large variation in phenotype, agamid lizards represent
a great system for comparative studies in multiple fields of
biology, including phylogeography (Macey et al., 1998; Mel-
ville et al., 2009; Moody, 1980), ethology (Kastle & Schleich,
1998; Murphy et al., 1978; Qi et al., 2011; Wei & Lin, 1981),
and evolutionary biology (Grismer, 2010; Schulte et al., 2002;
Stuart-Fox & Ord, 2004). Across mainland Asia, China har-
bors the greatest diversity of agamid lizards, currently pos-
sessing 52 currently recognized species in the country rep-
resenting four subfamilies (Ananjeva et al., 2011). Of this
diversity, nearly half (25 species) are believed to be endemic
to China, including majority of the enigmatic Mountain Drag-
ons of the genus Japalura Gray, 1853 (Ananjeva et al., 2011;
Wang et al., 2015). 1
To date, the genus Japalura is composed of 30 species, dis-
tributed across much of Asia (Wang et al., 2015). The vast
majority of this diversity occurs in the territory of Greater China
(16 species found in Mainland China, five species on the is-
lands of Taiwan; Ota et al., 1998; Wang et al., 2015). Until
recently, Japalura flaviceps Barbour & Dunn, 1919 was recog-
nized as having the widest geographic range in China, with
populations distributed from eastern Tibet to Gansu Province in
central China (Pope, 1935; Zhao et al., 1999). However, subse-
quent examinations of morphological variation among popula-
tions of J. flaviceps not only revealed the widespread species to
be a complex of distinct evolutionary lineages, but also re-
stricted the range of true J. flaviceps to the valleys of the Dadu
River in northwestern Sichuan only (Gao & Hou, 2002; Li et al.,
Received: 12 November 2015; Accepted: 10 December 2015
Foundation items: This work was supported by the Ministry of Science
and Technology of China (2014FY210200, 2011FY120200) and the
Animal Branch of the Germplasm Bank of Wild Species of Chinese
Academy of Sciences (the Large Research Infrastructure Funding).
*Corresponding author, E-mail: kai.wang-2@ou.edu; chej@mail.kiz.ac.cn
DOI: 10.13918/j.issn.2095-8137.2016.1.41

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42
2001; Manthe y et al., 2012; Wang et al., 2015). Further-
more, Manthey et al. (2012) and Wang et al. (2015) sug-
gested that unexamined populations of J. cf. flaviceps
outside of the Dadu River drainage may represent addi-
tional, unique evolutionary lineages within the species
complex, worthy of additional systematic study.
During herpetofaunal surveys of eastern Tibet in 2013 and
of northwestern Yunnan in 2015, we captured eight and 12
specimens of lizards in the upper Nujiang (=Salween) Valley,
eastern Tibet, China, and the upper Lancang (=Mekong)
Valley, northwestern Yunnan, China, respectively. All indi-
viduals were identified to the genus Japalura. Although
these populations are considered to be part of the J.
flaviceps species complex, they can be distinguished readily
from true J. flaviceps sensu Manthey et al. (2012) and other
members of the genus by suites of distinct morphological
characters. In this study, we describe these populations as
two distinct, new species within the genus. We review the
taxonomic history of the J. flaviceps species complex, com-
ment on the importance of coloration for species delimitation
in the genus and discuss directions for future taxonomic
studies of the group. Additionally, we assess the new spe-
cies and other imperiled congeners against the IUCN criteria
for classification (IUCN, 2013).
MATERIALS AND METHODS
Eight specimens of one new species were collected from the
upper Nujiang Valley near the Nujiang Bridge at Baxoi, Qamdo
Prefecture of eastern Tibet, China, including two adult males,
four adult females, and two juveniles. Twelve specimens of a
second new species were collected from the upper Lancang
Valley at Ninong, Deqin, northwestern Yunnan, China, including
11 adult males and one adult female.
Following euthanasia, tissue samples were taken from livers
and preserved in 95% ethanol, and voucher specimens were
fixed in 10% buffered formalin and later transferred to 70%
ethanol for long-term preservation. With the exception of a
single female specimen collected from Tibet that possesses an
incomplete tail (KIZ 014040), all adult specimens were chosen
as the type series. All specimens (including KIZ 014040) are
deposited in the Museum of Kunming Institute of Zoology, Chi-
nese Academy of Sciences (KIZ).
Measurements were made with digital calipers to the nearest
0.1 mm, except for snout-vent length (SVL) and tail length (TAL),
which was made with a ruler to the nearest 1 mm. With the
exception of several new traits measured in this study, focal
characters and character definitions follow Wang et al. (2015):
snout-vent length (SVL), tail length (TAL), head width (HW),
snout-eye length (SEL), fore-limb length (FLL), hind limb length
(HLL), supralabial count (SL), infralabial count (IL), middorsal
scale (MD), Toe IV subdigital lamellae (T4S), Toe IV length
(T4L), trunk length (TRL), interorbital distance (IOD), number of
scales between nasal and first supralabials (NSL), supraciliary
count (SCL), and number of scale rows between sixth su-
pralabial and orbit circle (SOR). Additionally, in this study we
examined the following morphometric characters (definitions
provided after colon): enlarged, conical, post-tympanic scale
count (PTY): large conical scales posterior to tympanum; and
enlarged, conical, post-rictal scale count (PRS): large conical
scales posterior to the rictus. Values for paired characters (SL,
IL, NSL, SOR) were recorded from both sides of the body, with
counts provided in left/right order.
Summaries of specimens examined are listed in Appendix
I. For comparisons, morphological data of the following
phenotypically similar species were collected from
vouchered specimens (from type or topotype specimens
when available): J. batangensis, J. dymondi, J. micang-
shanensis, J. flaviceps, J. splendida, J. varcoae, J. vela, J.
yunnanensis, and J. zhaoermii. Currently, with recognized
populations of J. splendida distributed across multiple dis-
tinct zoogeographic regions of China (Xie et al., 2004; Zhao
et al., 1999), and known to possess considerable variability
in morphological characters (Manthey et al., 2012; Yang &
Rao, 2008; Zhao et al., 1999), it is likely that the widespread
species represents a complex of unique evolutionary line-
ages. Therefore, to avoid unnecessary confusion in drawing
comparisons with potentially unique but undescribed diver-
sity, we include morphological data from the type specimen
of J. splendida as well as specimens from localities geo-
graphically proximate to the species’ type locality along the
Yangtze River (e.g. Chongqing). Morphological data of the
following species were obtained from the literature: J. brevi-
cauda (Manthey et al., 2012), J. kumaonensis (Schleich &
Kästle, 2002), J. luei (Ota et al., 1998), J. makii (Ota, 1989),
and J. yulongensis (Manthey et al., 2012).
Comparisons of coloration in life are based on type de-
scriptions and available color photographs (Manthey, 2010;
Yang & Rao, 2008; Zhao et al., 1999). Museum abbrevia-
tions for specimens examined follow Sabaj Perez (2015),
and include: Chengdu Institute of Biology, Chinese Academy
of Sciences (CIB); Kunming Institute of Zoology, Chinese
Academy of Sciences (KIZ); Museum of Comparative Zool-
ogy at Harvard University (MCZ), Boston, MA, USA; and
National Museum of Natural History (USNM), Washington
D.C., USA.
The topographic map shown in Figure 1 was created by N. A.
Huron in ArcMap v.10.3.1 using the digital elevation model
(DEM) layers based on NASA’s Shuttle Radar Topographic
Mission (SRTM). The SRTM data are available for free at ap-
proximately 90 meters resolution (3 arc-second projections;
Reuter et al., 2007; CIAT-CSI SRTM, 2015).
RESULTS
Japalura laeviventris sp. nov. Wang, Jiang, Siler, and Che
(Figures 1-4)
Synonyms
Japalura flaviceps Pope, 1935: 467; Zhao & Jiang, 1977: 293-298;
Hu et al., 1987: 112; Zhao et al., 1999: 111-115; Li et al., 2010: 115.
Holotype: KIZ 014038, adult male, collected near the Nujiang
Bridge in the upper Nujiang Valley at Baxoi (=Basu), Qamdo
(=Changdu), eastern Tibet (=Xizang), PR China (N30.10034,
E97.22787, 2 739 m elevation); collected by Ke JIANG on 3
July 2013.

Zoological Research 37(1): 41-56, 2016 43
Figure 1 Distribution map of Japalura in the Hengduan Mountain Range, southwest China (Map created by Nicholas A. HURON and Cameron D.
SILER)
Color-coded shapes show the distribution of type localities for the new species (stars), true J. flaviceps sensu Wang et al. (2015) (square), and other refer-
enced members of the J. flaviceps Species Complex (circles).
Paratopotypes: One adult male (KIZ 014037) and three adult
females (KIZ 014041-43); collected by Ke JIANG, Kai WANG,
and Ya-Qiang SUN.
Diagnosis: Following Inger’s definition of the genus (Inger,
1960), the new species is assigned to the genus Japalura
based on a number of diagnostic characters, including: (1)
dorsal scales unequal in size; (2) enlarged crest scales present;
(3) gular pouch present; (4) lateral fold of skin in axilla-groin
region present; (5) supraciliary scales greatly imbricate; (6)
head relatively long, flat; (7) tail long, slender; (8) tail cylindrical
in shape; and (9) precloacal and femoral pores absent.
Japalura laeviventris sp. nov. can be distinguished from all
congeners by the combination of the following suite of morpho-
logical characteristics: (1) small adult body size (SVL 67-72 mm
in males, 64-70 mm in females); (2) moderate TAL (TAL/SVL
168%-200%); (3) moderate HLL (HLL/SVL 64.3%-78.4%); (4)
NSL 1; (5) T4S 22-26; (6) SOR 3; (7) strongly-protuberant,
conical, post-tympanic scale absent; (8) strongly-protuberant,
conical, post-rictal scale absent; (9) tympanum concealed; (10)
nuchal crests relatively raised on weak skin folds; (11) dorsal
crests weakly developed without distinct skin folds in males; (12)
transverse gular fold present; (13) gular pouch distinct, present;
(14) scales of ventral surface of body smooth or weakly keeled;
(15) MD 57-59, (16) ground dorsal coloration off-white in males,
brownish-gray in females; (17) dorsal, lateral, and ventral surface
of head, dorsal forelimbs, and lateral surface of body speckled
with black; (18) distinct radial streaks around eyes; (19) dorso-
lateral stripes present, smooth-edged, pale-yellow in males; (20)
dark-brown, “M”-shaped pigmentation patterns along dorsal
midline in males; and (21) small, triangular, orange gular spots in
adults of both sexes.
Description of holotype: Adult male, SVL 67 mm, TAL 133 mm,

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FLL 30.6 mm, HLL 49.8 mm, HW 15.5 mm, HL 21.3 mm. Rostral
rectangular, three times broader than high, in contact with six
small scales excluding supralabial. Nasal sub-circular, bordered
by 10/9 small scales. Single scale between rostral and first su-
pralabial. Supralabials 7/8, smooth, posteriormost longest. Loreal
scales irregularly arranged and weakly keeled. Ciliaries circular,
much smaller than other dorsal scales, forming orbit circle; su-
praciliaries 9/11, prominent, elongated, above orbit; first 5/7 su-
praciliaries overlapping one-half to two-third of its length with sub-
sequent ones, last four slightly overlapping; three rows of scales
between orbit circle and sixth supralabials on both sides of head, all
weakly keeled, with scales of middle row largest. Scales posterior
to eyes strongly keeled; five enlarged scales between orbit and
tympanum on both sides; tympanum covered with much smaller
scales. Dorsal head scales heterogeneous in size, approximately
circular shaped, mildly keeled, convex, somewhat granular in
appearance; single row of five scales forming a weak ridge along
snout midline from one scale posterior rostral to mid-point between
anterior corner of eyes; parietal smooth, enlarged, with a distinct
pineal eyespot; post-occipital and posterior lateral head scales
strongly keeled; conical scale 1/0 on occipital region, weakly devel-
oped; no large conical scales posterior to tympanum or rictus.
Ventral head scales homogeneous in size, mostly smooth,
weakly keeled posterolaterally; transverse gular fold distinct, well
developed; gular pouch present; shoulder fold posterior to gular
fold on each side, from ventral surface of neck to just about 5 mm
above pectoral joint; axillary fold present on each side of body.
Middorsal crest scales 59, roughly equal in size to neighboring
scales, imbricate along body midline; nuchal crest relatively
raised on skin fold, while dorsal crest weakly developed without
skin folds. A row of enlarged and distinctively keeled scales run-
ning parallel to dorsal crest from pectoral region of body to pelvis on
each side of crest above dorsolateral stripes. Ground dorsal scales
heterogeneous in size; scales of axilla much smaller than remain-
ing dorsals; large, distinct, flat scales distributed irregularly across
dorsal surface of body, circular in shape, roughly four times larger
than ground scales, at times arranged in proximally transverse
rows. Dorsal limb scales distinctively keeled, roughly equal in size
to ventral scales, homogeneous in size on forelimbs, heterogene-
ous on hind limbs; Toe IV subdigital lamellae 25/24. Tail scales
keeled in lateral rows; cloaca scales small. Ventral scales of body
and limbs mostly homogeneous in size, smooth or weakly keeled
close to lateral sides.
Coloration of holotype in life: In life, the dorsal surface of
the head is light yellowish, speckled with small black scales,
of which some connect to form dark X-shaped patterns. Two
of these X-shaped patterns are observed on the snout, and
another can be observed on the dorsal surface of the head
between the orbits. The lateral surface of the head is off-white
and is also speckled with black scales. Some black scales
form radial stripes around the eyes, with the stripes directed
posteriorly being the broadest. The ground coloration of the
dorsal and lateral surfaces of the body is off-white. Smooth-
edged, pale-yellow dorsolateral stripes run along the entire
length of the body on each side of the vertebral crest. Black
vermiculate pigmentation patterns are present on the lateral
surfaces of the body, beneath and proximate to the dorso-
lateral stripes. A series of three V-shaped brownish-black
patterns of dark pigmentations are observed on the dorsal
surface of the body, distributed from the occipital region of the
head to the pectoral joint of the body along the dorsal midline.
These shapes are followed posteriorly by a series of four
brownish-black, M-shaped pigmentations, of which the last
one is gradually faded and somewhat unclear. Numerous
brownish-gray, transverse bands are present on the posteri-
ormost two-thirds of the tail. The bands do not form complete
rings around the tail ventrally. The dorsal surfaces of the fore-
limbs are off-white and speckled with black scales. The black
scales in this region are connected in places by gray-blotched
scales, and together, these colors create irregular transverse
bands of dark coloration. The dorsal surfaces of the hind
limbs are light gray, with dark-gray transverse patterns similar
to those observed on the fore-limbs. The ground coloration of
the ventral surface of the head is uniform white, but heavily
speckled with black scales. Sporadically, small series of these
black scales from short, longitudinal, vermiculate stripes. A
triangular, bright orange gular spot is present in the center of
the gular pouch. The ventral surfaces of the body, limbs, and
tail are uniform white, except for the ventral surfaces of the
hands and feet, which are more yellowish (Figure 2).
Coloration of holotype in preservative: The coloration of the
holotype in preservative closely matches its coloration in life; how-
ever, the following differences are observed: (1) the ground
coloration of the dorsal surface of the head as well as the col-
oration of the dorsolateral stripes is light gray, and (2) the trian-
gular orange gular spot turns white (Figure 3).
Variation and sexual dimorphism: Variations in pholidosis
and measurements are summarized in Table 1. The male para-
topotype (KIZ 014037) is slightly darker in dorsal coloration than
the holotype in preservative. Females are sexually dimorphic
from males by having shorter snouts (SEL/HL), less speckled
ventral surfaces of the heads (v.s. heavily speckled), more
consistently arranged transverse rows of enlarged scales on
the dorsal surfaces of the body (v.s . irregularly distributed),
smaller orange gular spots (v.s . larger), much more distinct
transverse patterns of pigmentations at bases of the tails (v.s .
less distinct), as well as by the absences of M-shaped dark
pigmentation patterns along the dorsal midline of the body (v. s.
presence), and the presence of dark transverse bands on the
dorsal surface of the body that extend to the lateral surfaces of
the body (v.s. absence). Yellow dorsolateral stripes are absent
in most females, except for a single individual (KIZ 014043),
which has wavy dorsolateral stripes. Juveniles (KIZ 014039,
014044) closely resemble adult females, except for the ab-
sence of orange gular spots. A single juvenile (KIZ 014039) was
observed to possess a light-gray, dorsal, vertebral stripe from
the pectoral region of the body to pelvis. This trait was not
observed in any adult females.
Comparisons: Populations of the new species were identified
previously as J. flaviceps. However, the new species can be

Zoological Research 37(1): 41-56, 2016 45
Figure 2 Lateral and ventral views of adult male holotype KIZ 014038 (A and B) and adult female paratopotype KIZ 014043 (C and D) of Ja-
palura laeviventris sp. nov. in life (Photos by Kai WANG)
Note the dark reddish orange color posterior to the shoulder fold in the lateral view is the color of ectoparasites and not a coloration pattern of the new species.
Figure 3 Dorsolateral views and ventral close-ups views of adult male holotype KIZ 014038 (A and B) and adult female paratopotype KIZ
014043 (C and D) of Japalura laeviventris sp. nov. in preservative (Photos by Kai WANG)
distinguished readily from the latter by having smooth or weakly
keeled scales on the ventral surface of the head and the body
(v.s . distinctively keeled), a greater number of MD (57-59 v.s .
43-48), heavily speckled ventral surfaces of the head, with
speckles rarely forming short lines (v. s. speckles absent, but
broad, dark stripes present and interconnected into a mosaic
pattern), as well as by the absence of strongly-protuberant,
conical, post-rictal scale (v.s . presence), presence of X-shaped
patterns of dark pigmentation on the dorsal surface of the head
(v.s . absence, or presence of few transverse streaks), presence
of distinct radial patterns around the eyes (v. s. absence), pres-
ence of M-shaped patterns of dark pigmentation along the
dorsal midline between the two dorsolateral stripes in males (v. s.
rhombus-shaped patterns with distinct yellow centers), and
presence of distinct orange gular spots in both sexes (v.s . ab-
sence in both sexes).

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Zoological Research 37(1): 41-56, 2016 51
Japalura laeviventris sp. nov. is most similar to J. kumaonen-
sis (Annandale, 1907) and J. yunnanensis Anderson, 1879 in
coloration patterns. All three species possess radial patterns of
dark pigmentation around the eyes and light dorsolateral
stripes in males. However, the new species can be distin-
guished readily from the latter two by having smooth or
weakly keeled scales on the ventral surfaces of the head and
the body (v.s. distinctively keeled), a greater number of MD
(≥57 v.s. ≤52), as we as by the absence of strongly protuber-
ant, conical, post-tympanic scales (presence and in high num-
bers) and absence of strongly-protuberant, conical, post-rictal scale
(v.s . presence). Additionally, Japalura laeviventris sp. nov. differs
from from J. kumaonensis by having a greater number of SL (7-
9 v. s. 5 or 6), a concealed tympanum (v.s . exposed), a relatively
well developed gular pouch (v.s . weakly developed), M-shaped
patterns along the dorsal-midline in males (v.s. chevron-shaped),
the posteriorly directed radial-stripes of the eyes less prominent
and short, ending before reaching the tympanums (v.s . distinct,
broad, enclosing the tympanums), as well as by the presence of
a transverse gular fold (v. s. absence) and presence of orange
gular spots in both sexes (v.s. absence in both sexes); and from
J. yunnanensis by having a shorter tail (TAL/SVL ≤200% v.s.
≥235%), fewer T4S (22-26 v.s. 27-31), greater number of NSL
(1 v.s . 0), broad dorsolateral stripes with smooth edges in males
(v.s . narrow and jagged), an off-white ground coloration on the
dorsal surface of the body (v.s . green or brown), the terrestrial
lifestyle (v.s . arboreal), as well as by the presence of a trans-
verse gular fold (v.s . absence) and presence of orange gular
spots in both sexes (v.s. light yellow gular spots in males, some-
times absence in females).
In addition to the four species compared above, the new spe-
cies can be diagnosed from all remaining congeners by
having smooth or weakly keeled scales on the ventral
surfaces of the head and the body (v. s . distinctively
keeled), an off-white ground coloration of the body (brown,
black, or green), heavily speckled surfaces of the head and
lateral surfaces of the body (v. s . absence or weakly speck-
led lateral body only), M-shaped dark patterns of pigmenta-
tions along the dorsal midline of the body between the two
dorsolateral stripes in males (v.s. rectangular blotches of
dark pigmentations), and the orange gular spots in both
sexes in life (v.s. other colorations, in males only), as well
as by the absence of large, conical post-rictal and post-
tympanic scales (v.s . presence and in high numbers).
Distribution and Natural History: Japalura laeviventris sp.
nov. is known only from its type locality in the upper Nujiang
Valley (Figure 1); however, the species may also occur
along geographically proximate stretches of Nujiang. Little
vegetation exists in the Hot-and-Dry Valley (Figure 4). The
new species is terrestrial, with individuals have been ob-
served on the sandy hills alongside the river, utilizing rock
crevices and rodent burrows as shelters. No larger reptiles
have been recorded from the area. Males of the new spe-
cies are more conspicuous in behavior than females, and
can be observed basking for greater lengths of time in open
habitat. In contrast, females appear to stay in closer prox-
imity to shelters.
Figure 4 Preferred microhabitat of Japalura laeviventris sp. nov.
near the Nujiang Bridge, Baxoi County, Qamdo Prefecture, east-
ern Tibet, China (Photo by Ya-Qiang SUN)
Etymology: We derive the new species name from the Latin
word “laeviventris, ” meaning “smooth venter, ” in reference to
one of the major diagnostic characteristics of the new species:
smooth or weakly keeled ventral body scales. Suggested com-
mon name: Smooth-venter Mountain Dragon (English); Hua Fu
Pan Xi (Chinese; 滑腹攀蜥).
Japalura iadina sp. nov. Wang, Jiang, Siler, and Che
(Figures 5-7)
Synonyms
J. flaviceps Zhao et al., 1999: 293-298; Yang and Rao, 200:
200-201; 8; Xu and Zhang 2011: 202-203; J. splendida Xu and
Zhang 2011: 202-203; J. cf. flaviceps Manthey et al., 2012
Holotype: KIZ 019321, adult male, collected by Kai WANG on
27 May 2015, from the Lancang Valley at Ninong, Deqin,
northwest Yunnan, China (N28.370255, E98.865287, 2 062 m
elevation).
Allotopotype: KIZ 09398, adult female. Collected by Da-Hu
ZOU. Specimen shares the same locality and collection infor-
mation as the holotype.
Paratopotypes: KIZ 09401-03, 019322, 019323, 019325-28, all
adult males. Collected by Kai WANG, Ke JIANG, and Da-Hu
ZOU.
Diagnosis: Following Inger’s (1960) definition of the genus, the
new species is assigned to Japalura based on a number of
diagnostic characters, including: (1) dorsal scales unequal in
size; (2) enlarged crest scales present; (3) gular pouch pre-
sent; (4) lateral fold of skin in axilla–groin region present; (5)
supraciliary scales greatly imbricate; (6) head relatively long,
flat; (7) tail long, slender; (8) tail cylindrical in shape; and (9)
precloacal and femoral pores absent.
The new species differs from all congeners by a combination
of the following morphological characters: (1) gular fold present;
(2) distinct gular pouch present; (3) relative hind-limb length
moderate HLL/SVL 69.6%-80.1%; (4) relative tail length

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Figure 5 Dorsolateral (A), ventral (B), and ventral head close-up
views (C) of the adult male holotype (KIZ 019321) of Japalura
iadina sp. nov. in life (Photos by Kai WANG)
moderate TAL/SVL 173%-198%; (5) F4S 15-17; (6) T4S 19-25;
(7) MD 35-46; (8) three lateral rows of enlarged scales pre-
sent on the dorsal surface of the body parallel to the dorsal
crest; (9) nuchal and dorsal crests moderately raised on skin
folds; (10) nuchal and dorsal crests relatively low and discon-
tinuous in males; (11) dorsal and lateral surfaces of body
emerald green in males, yellowish brown speckled with large,
light yellow scales in females; (12) tail yellowish green in males,
brownish in females; (13) ventral surface of body bluish or
whitish gray, sometimes with black speckles, in males, uniform
yellow in females; (14) two smooth-edged, light-green, dorso-
lateral stripes present in males; (15) vermiculate stripes on
ventral surface of head present, distinct, blackish blue in males,
black in females; (16) gular region dark blue in males, yellow in
females.
Description of holotype: Adult male, SVL 60 mm, TAL 116 mm,
TrL 28.3 mm, HL 18.5 mm, HW 12.4 mm, IOD 9.1 mm, SEL 6.9 mm,
FLL 28.8 mm, HLL 44.3 mm, T4L 10.3 mm. Rostral rectangular,
three times broader than high, in contact with six small scales
excluding supralabial; nasal sub-circular; single scale between
Figure 6 Dorsolateral (A), ventral (B), and ventral head close-up
views (C) of the adult female allotype of Japalura iadina sp. nov.
(KIZ 09398) in life (Photos by Kai WANG)
Figure 7 Habitat of Japalura iadina sp. nov. in the Dry-Hot Valley
of Lancang River, Deqin, Northwest Yunnan, China (Photo by Kai
WANG )
nasal and first supralabial; supralabials eight on both sides of
head, weakly keeled; loreal scales irregularly arranged and
moderatly keeled; ciliaries circular, much smaller than other
scales of lateral head, forming orbit circle; supraciliaries six on
both sides of head, prominent, elongated; three rows of scales
between orbit circle and sixth supralabial on both sides of head,
all weakly keeled; scales posterior to eyes strongly keeled; orbit

Zoological Research 37(1): 41-56, 2016 53
and tympanum separated by five enlarged scales on both sides
of head; tympanum covered with small scales; dorsal head
scales heterogeneous in size, distinctively keeled, convex; single
row of five scales forming weak ridge along snout midline from
just posterior to rostral to point in line with anterior corner of eyes;
parietal keeled, enlarged; pineal eyespot present; post-occipital
and posterior lateral head scales strongly keeled; conical scales
on occipital region numerous, strongly protuberant; conical scales
posterior to tympanum two, large; conical scale posterior to tym-
panum or rictus single, large, on each side of head.
Ventral head scales homogeneous in size, distinctively
keeled; transverse gular fold present, distinct; gular pouch
present; shoulder fold posterior to gular fold on each side of
body present, distributed from ventral surface of throat to dorso-
lateral stripes; axillary fold present on each side of body. Mid-
dorsal scales 40, larger than neighboring scales, imbricate
along dorsal midline; nuchal and dorsal crests slightly raised on
skin folds, with distinct break separating regional folds; three
dorsolateral rows of enlarged, distinctively keeled scales run-
ning parallel to dorsal crest from pectoral region of body to
pelvis on each side of body, with one dorsal to dorsolateral
stripe and two along upper and lower edges of dorsolateral
stripe; ground dorsal scales heterogeneous in size; scales of
axilla smaller than dorsal scales; large conical scales present,
on dorsal and lateral surface of body, distinctly keeled, ran-
domly scattered; dorsal limb scales distinctively keeled, slightly
larger than ventral scales, homogeneous in size on fore-limbs,
heterogeneous on hind limbs; F4S 16/16; T4S 20/21. Tail
scales keeled, in lateral rows; cloaca scales small. Ventral
scales of body and limbs near homogeneous in size, distinc-
tively keeled.
Coloration of holotype in life: The ground coloration of the
dorsal, lateral, and ventral surfaces of the head is emerald
green. Four broad, black transverse bands are observed on the
dorsal surface of the head, and are equally spaced across the
region between the nares and a point in line with the posterior
edge of the orbits. Black reticulated patterns are observed
posterior to the last transverse bands on the dorsal surface of
the head. Nine black streaks are observed radiating around the
eye on each side of the head. The streaks extend ventrally to
the supralabial scales, with the posteriormost streaks broadest.
The infralabial scales and the margin of the lower jaw are em-
erald green with black bars present on several of the infralabial
scales. These infralabial bars match up with streaks present on
the supralabial scales, and extend extend further posteroven-
trally, gradually transitioning into blackish blue, vermiculate
stripes on the ventral surface of the head. The vermiculate
stripes connect posteriorly with the large, triangular shaped,
blue gular spot located in the center of the gular pouch.
A black vertebral stripe runs from the occipital region of the
head to the pelvis along the dorsal midline of the body. Several
crest scales and a few small, ground scales along the vertebral
stripe are green. The lateral and dorsolateral surfaces of the
body are emerald green, with reticulated patterns of black pig-
mentation on the lateral surfaces of the body. These reticulated
patterns of pigmentation form a thin, black, dorsolateral line
along the ventral edge of the dorsolateral stripe on each side of
the body. The dorsolateral stripes are green and smooth-edged,
running from the posterior occipital region of the head to the
pelvis. A series of enlarged, distinctively keeled, green scales
are observed in a dorsolateral series on each side of the dorsal
crest, running from the neck to the pelvis and distributed along
the black vertebral stripe. The ventral surface of the body is
uniform whitish blue and is slightly lighter anteriorly.
The ground coloration of the dorsal surfaces of the limbs is
emerald green, with numerous transverse bands running from
the proximal to distal regions of the limbs. The ventral surfaces
of the fore-limbs and hind limbs are whitish blue and greenish
yellow, respectively. The ventral surfaces of the hands and feet
are gray in coloration.
The dorsal surface of the tail is greenish yellow, with numer-
ous dark gray, transverse bands running along its length. The
transverse bands do not form complete rings around the tail
ventrally. The ventral side of the tail is significantly duller gray in
coloration (Figure 5).
Coloration of holotype in preservative: The coloration of the
holotype in preservative closely resembles its coloration in life;
however, the following differences are observed: 1) the emerald
green coloration on some parts of the dorsal surfaces of the
head, body, and limbs changed to light blue, and 2) the blue
coloration of the gular spot on the ventral surface of the head
and body faded significantly.
Variation and sexual dimorphism: Variation in morphometric
characters and pholidosis patterns is summarized in Table 2. The
new species is sexually dimorphic, with females possessing
distinct coloration and pigmentation patterns from males. The
female allotype of the new species differs from males by having
light, yellowish brown ground coloration on the dorsal surfaces of
the head, body, and limbs (v.s . emerald green), distinct, light
brown transverse bands on the dorsal surface of the body (v.s .
single, black vertebral stripe), a greenish yellow gular spot (v. s.
blue), and a white ground coloration on the ventral surfaces of the
body and limbs (v.s . blue; Figure 6).
Comparisons: The new species Japalura iadina sp. nov. has
been confused previously with J. flaviceps; however, it can be
distinguished readily from the latter by having a smaller adult
body size (SVL 54-65 mm v.s. 64-75 mm), a tendency to-
wards a shorter relative snout length (SEL/HL 34.9%-40.2%
v.s . 40.0%-44.2%), smaller, less protuberant nuchal and
dorsal skin folds in males (v.s. strongly protuberant), distinct
ground coloration on the dorsal surfaces of the head, limbs,
and body in males (emerald green v.s. brown), and distinct
patterns of pigmentation along the dorsal midline of the body
in males (vertebral black stripes v.s. a series of dark rhom-
boid-shaped patterns of pigmentation), as well as by the
presence of distinct gular spots in both sexes (v. s. absence),
and by the presence of distinct black radial stripes around
the eyes (v.s . absence).
The new species is morphologically most similar to J. splen-
dida, J. makii, and J. luei, of which all four species have the green

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54
background coloration. However, Japalura iadina sp. nov. can be
diagnosed from J. splendida by having a terrestrial life style (v.s.
arboreal), a smaller adult body size (SVL 54-65 mm v.s . 59-92
mm), fewer T4S (19-24 v.s. 24-28), a shorter relative tail length
(TAL/SVL≤198% v.s. ≥217%), distinct tail coloration in males
(greenish yellow v.s . brownish gray), as well as by the absence
of white lip stripes (v.s . presence), presence of gular spots in
both sexes (v. s. absence), and absence of green coloration on
the dorsal surfaces of the body in females (v.s . presence); from
J. makii by having a smaller adult body size (SVL 54-65 mm v.s .
61-79 mm), a relatively shorter tail (TAL/SVL 1.73-1.98 v.s .
2.09-2.61), fewer T4S (19-24 v.s. 24-30), a pink coloration of
tongue and oral cavity in life (v. s. yellow), a terrestrial lifestyle
(vs. arboreal), as well as by the absence of a gular fold (v.s.
presence), absence of green, transverse bands across the
dorsal surface of body (v.s. presence), and absence of lip
strips below eyes (v.s. presence); and from J. luei by having a
smaller adult body size (SVL 54-65 mm v.s . 65-75 mm), a
relatively shorter tail (TAL/SVL 1.73-1.98 v.s . 2.31-2.48), fewer
T4S (19-24 v. s. 24-28), a terrestrial lifestyle (v. s. arboreal),
continuous dorsolateral stripes in males (v. s. a dorsolateral
series of disconnected, irregular, large markings), as well as by
the absence of a gular fold (v. s. presence), and absence of lip
stripes below eyes (v.s . presence).
Japalura iadina sp. nov. differs from J. laeviventris by having
a smaller adult body size (SVL 54-65 mm v.s . 64-72 mm), dis-
tinctively keeled ventral scales of head and body (v.s. smooth or
weakly keeled), fewer MD (35-46 v. s. 57-59), distinct ground
coloration of the dorsal surfaces of head and body in males
(emerald green v. s. off-white), distinct coloration of gular spots
(blue in males, greenish yellow in females v.s . orange in both
sexes), and distinct patterns of pigmentations on the dorsal
surfaces of the body along the dorsal midline (black vertebral
stripes speckled with green v. s. M-shaped patterns of dark
brown pigmentation).
Additionally, when compared with species distributed in close
geographic proximity along the Lancang River, J. iadina sp. nov.
can be distinguished from J. vela by having less protuberant,
discontinuous vertebral crests in males (v. s. sail-like, strongly
protuberant, continuous), distinct ground coloration on the
dorsal surfaces of the body (emerald green v.s . black) and tail
(yellow v.s . gray) in males, uniquely shaped dorsolateral stripes
in males (smooth edged v.s. strongly jagged), as well as by the
presence of gular spots in both sexes (v. s. absence in both
sexes), and by the absence of weakly defined, reddish dorso-
lateral lines in females (v.s . presence); and from J. yunnanensis
by having a terrestrial lifestyle (v.s . arboreal), shorter relative tail
length (TAL/SVL ≤198% v.s.≥237%), fewer T4S (19-24 v.s .
27-31), distinct coloration of gular spots (blue in males, greenish
yellow in females vs. yellow in both sexes when present or
absence in females), as well as by the presence of a transverse
gular fold (v.s . absence).
Distribution and natural history: Although locally abundent,
Japalura iadina sp. nov. is known only from the type locality
(Figure 1). The new species is terrestrial, inhabiting dry, rocky
habitats along the Lancang River (Figure 7). The emerald green
coloration of the males makes them stand out from the rocky,
environmental background. Tail autotomy has been observed
for this population.
Etymology: The Latin name “iadina” means “emerald like, ”
which describes the diagnostic emerald green body coloration of
males of the new species. Suggested common name: Emerald
Mountain Dragon (English), Fei Cui Pan Xi (Chinese; 翡翠攀蜥).
DISCUSSION
Body coloration, particularly the coloration of the gular region,
has been suggested to play an important role in the recognition
of conspecifics, sexual selection, and the general diversification
of agamid lizards in general (Bastiaans et al., 2014; LeBas &
Marshall, 2000; Stuart-Fox & Ord, 2004), particularly members
of the genus Japalura (Kästle & Schleich, 1998; Wei & Lin,
1981). Within the genus Japalura, color patterns vary among
species from mainland Asia, but appear to be conservative within
species, and hence serve as good diagnostic characters for
systematic studies of species diversity (Wang et al., 2015). How-
ever, to date, detailed descriptions of coloration in life, have not
been published for many species in the genus, including J. brevi-
cauda, J. chapaensis, J. grahami, J. hamptoni, J. otai, J. yulon-
gensis; (Mahony, 2010; Manthey, 2010; Manthey et al., 2012; Ota
& Weidenhöfer, 1992; Zhao et al., 1999). Since specimen colora-
tion is recognized to fade significantly after preservation for many
specimens in natural history collections, color descriptions of
preserved specimens may not be as applicable to diagnostic
comparisons. Therefore, we recommend future research focusing
on describing coloration patterns based on new observations of
species of Japalura in life.
Despite the importance of the coloration patterns in the taxo-
nomic studies of members of the genus Japalura, the diagnos-
tic coloration of J. flaviceps has been historically confused,
which has continued to the species’ recognized wide spread
distribution. Originally described by Barbour & Dunn (1919)
from the Tung River Valley (today Dadu River Valley) in north-
west Sichuan Province, J. flaviceps was described as possess-
ing a uniform, “dusky brown” dorsal head coloration and lacked
“strongly marked stripe from the eye to the angle of the mouth.”
Together with photographs of the holotype of J. flaviceps (MCZ
R-12469) and topotypic males from the Dadu River Valley,
Manthey et al. (2012) argued that J. flaviceps did not possess
dorsal and lateral pigmentation patterns on the head, and pro-
posed that the absence of such patterns should be used as a
diagnostic feature of the species. Although most topotypic
specimens of J. flaviceps that we examined possess no distinct,
radial pigmentation patterns around the eyes, we did observe
several individuals, particularly subadults, that possess several
distinct transverse bands of darker pigmentation across the
dorsal region of the head (CIB 2333 and 2549). Therefore, we
suggest that the absence of radial patterns of darker pigmenta-
tion around the eyes may be a more consistent feature of J.
flaviceps for use in diagnostic comparison. Furthermore, we
propose that the diagnostic feature of the true J. flaviceps should
be restricted to the following combination of morphological char-

Zoological Research 37(1): 41-56, 2016 55
acters: SVL 70-83 mm in adult males, 58-78 mm in adult fe-
males; TAL/SVL 165%-192%; HLL/SVL 62%-78%; SEL/HL
39%-43%; SOR 4; SL 9-11; IL 9-11; MD 43-54; T4S 21-24;
tympanum concealed; distinct gular fold present; distinct gular
pouch present; enlarged, conical, post-rictal, post-occipital, and
post-tympanic scales present, numerous, prominent; ventral
scales distinctively keeled; dorsal ground coloration of head and
body brownish gray; distinct radial stripes around eyes absent;
dark, broad, interconnected, vermiculated stripes on the ventral
surface of head present; gular spot absent; smooth-edged,
yellow dorsolateral stripes in males present; a series of rhomb-
shaped patterns of dark brown pigmentation with distinct yellow
centers present along the dorsal midline of the body.
Due to the continued taxonomic confusion over comparisons
of populations of the J. flaviceps species complex, J. flaviceps
was thought to be a widespread species, and its conservation
status was assessed as Least Concern (IUCN, 2013). However,
because of the absence of data allowing for confirmation of
species level diversity within the J. flaviceps Species Complex,
and the paucity of available information about intraspecific
ecological and genetic diversity, we recommend that J.
flaviceps and all conspecifics in the species complex be con-
sidered Data Deficient, and recommended immediate research
be focused on better understanding this unique complex of
agamid lizards.
Unfortunately, habitats in the Hengduan Mountain Range are
experiencing currently considerable human-mediated modifica-
tions as a result of the rapid development of hydropower plants
(Pan et al., 2002; Chen & Rao, 2010; personal communication
with Mr. Li CHENG). Although populations of Japalura can be
abundant, continued alteration of these valley habitats could
result in the extirpation of micro-endemic populations, or worse,
unique evolutionary lineages not yet recognized formally as
species. Additionally, many species of Japalura are experienc-
ing over-exploitation through the illegal pet-trade, both domesti-
cally and internationally (personal communication with Mr. Jia-
Wei WU and Mr. Mian HOU). Therefore, it is critical that re-
searchers and conservation and government agencies work
together to assess the conservation statuses and ecological
requirements of species in the genus, particularly species and
populations endemic to the river valleys in the range of the
Hengduan Mountain Range.
ACKNOWLEDGEMENTS
We would like to thank Dr. Gernot Vogel (Society for Southeast Herpetology,
Germany), Mr. Xin-Lei ZHAO (Chinese Academy of Medical Sciences), Mr.
Scott Farnsworth (Washington State University), and Dr. Su LIU (Shanghai
Chenshan Botanical Garden) for providing suggestions on the scientific
names of the new species, Mr. Duan YOU and Mr. Ya-Qiang SUN for assist-
ing our fieldwork in Tibet, 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 letting us examine specimens under their care, Mr.
Nicholas A. Huron (University of Oklahoma) for assistance with ArcGIS and in
the creation of the Figure 1 map, Ms. Jing-Ting LIU (Washington State Uni-
versity) and Ms. Dan-Lin LI (University of Idaho) for editing the photos, Mr.
Mian HOU for providing valuable information, and members of the Siler Lab
for their generous comments on the manuscript.
REFERENCES
Ananjeva N, Guo XG, Wang YZ. 2011. Taxonomic diversity of Agamid
lizards (Reptilia, Sauria, Acrodonta) from China: a comparative analysis.
Asian Herpetological Research, 2 (3): 117-128.
Anderson J. 1879 “1878”. Reptilia and Amphibia. In: Anatomical and Zoo-
logical Researches: Comprising an Account of the Zoological Results of the
Two Expeditions to Western Yunnan in 1868 and 1875. London: Bernard
Quarich, 803-804.
Annandale N. 1907. Reptiles. In: Boulenger GA, Annandale N, Wall F,
Regan CT. Report on a collection of Batrachia, Reptiles and Fish from
Nepal and the Western Himalayas. Records of Indian Museum, 1: 149-158.
Barbour T, Dunn ER. 1919. Two new Chinese Japaluras. Proceeding of the
New England Zoological Club, 7: 15-19.
Bastiaans E, Bastiaans MJ, Morinaga G, Gaytán JGC, Marshall JC, Bane B,
Cruz FM, Sinervo B. 2014. Female preference for sympatric vs. allopatric
male throat color morphs in the mesquite lizard (Sceloporus grammicus)
species complex. PLOS ONE, 9(4), e93197. doi: 10.1371/journal.pone.
0093197.
Chen XL, Rao DQ. 2010. Investigation of fish, amphibian, and reptile
species in the Shangri-La County and areas around small Zhongdian Key
Water Control Project. Journal of Yunnan Agricultural University, 25(4):
535-541. (in Chinese)
CIAT-CSI SRTM. 2015. The CGIAR Consortium for Spatial Information.
Available from: http: //srtm.csi.cgiar.org (1 December 2015).
Gao ZF, Hou M. 2002. Description of a new Japalura species from western
Sichuan Province, China. Sichuan Journal of Zoology, 21 (1): 3-5. (in Chinese)
Gray JE. 1853. Descriptions of some undescribed species of reptiles
collected by Dr. Joseph Hooker in the Khassia Mountains, East Bengal, and
Sikkim Himalaya. Annals and Magazine Natural History, 12 (2): 386-392.
Grismer JL. 2010. Phylogeny, taxonomy, biogeography and evolution of
asexuality within the southeast Asian lizard genus Leiolepis Cuvier, (1829).
Master Thesis. Villanova University, 1–70.
Hu S, Zhao E, Jiang Y, Fei L, Ye C, Hu Q, Huang Q, Huang Y, Tian W. 1987.
Amphibia-Reptilia of Xizang. Beijing: Science Press, 1-153. (in Chinese)
Inger RF. 1960. A review of the Agamid lizards of the genus Phoxophrys
Hubrecht. Copeia, 1960(3): 221-225.
IUCN. 2013. IUCN Red List of Threatened Species. Version 2013.2, Avail-
able from: http: //www.iucnredlist.org/ (10 November 2015).
Kästle W, Schleich HH. 1998 Studies on the Systematics and Biology of the
Genus Japalura (Sauria: Agamidae). Notes on Comparative Ethology and
Taxonomy of the Genus Japalura. In: Kästle W, Schleich HH (Ed.). The
Contributions to The Herpetology of South Asia (Nepal, India), Fuhlrott
Museum, Brand 4 (1998): 233-246.
LeBas NR, Marshall NJ. 2000. The role of colour in signalling and male
choice in the agamid lizard Ctenophorus ornatus. Proceedings of the Royal
Society of London B: Biological Sciences, 267 (1442): 445-452.
Li C, Deng QX, Wu Y, Wang Y. 2001. A new species of Japalura from
Sichuan (Agamidae Gray. Japalura). Journal of Sichuan Teachers College
(Natural Sciences), 22 (4): 329-331. (in Chinese)
Li PP, Zhao EM, Dong BJ. 2010. Amphibians and Reptiles of Tibet. Beijing:
Science Press, 1-251. (in Chinese)
Macey, JR, Schulte JA, Ananjeva NB, Larson A, Rastegar-Pouyani N,
Shammakov SM, Papenfuss TJ. 1998. Phylogenetic relationships among

www.zoores.ac.cn
56
Agamid lizards of the Laudakia caucasia species group: testing hypotheses
of biogeographic fragmentation and an area cladogram for the Iranian
Plateau. Molecular Phylogenetics and Evolution, 10 (1): 118-131.
Mahony S. 2010. Systematic and taxonomic revaluation of four little
known Asian agamid species, Calotes kingdonwardi Smith, 1935, Ja-
palura kaulbacki Smith, 1937, Salea kakhienensis Anderson, 1879 and
the monotypic genus Mictopholis Smith, 1935 (Reptilia: Agamidae).
Zootaxa, 7(2514): 1-23.
Manthey U. 2010. Agamid Lizards of Southern Asia – Draconinae 2,
Leiolepidinae. Chimaira, Frankfurt/M, 1-168.
Manthey U, Wolfgang D, Hou M, Wang XH. 2012. Discovered in historical
collections: two new Japalura species (Squamata: Sauria: Agamidae) from
Yulong Snow Mountains, Lijiang Prefecture, Yunnan, PR China. Zootaxa,
30(1664): 27-48.
Melville J, Hale J, Mantziou G, Ananjeva NB, Milto K, Clemann N. 2009.
Historical biogeography, phylogenetic relationships and intraspecific diver-
sity of agamid lizards in the central Asian deserts of Kazakhstan and
Uzbekistan. Molecular Phylogenetics and Evolution, 53(1): 99-112.
Moody S. 1980. Phylogenetic and historical biogeographic relationships of
the genera in the Agamidae (Reptilia, Lacertilia). PhD Dissertation, Univer-
sity of Michigan, 1–370.
Murphy JB, Lamoreaux WE, Carpenter CC. 1978. Threatening behavior in
the angle-headed dragon, Goniocephalus dilophus (Reptilia, Lacertilia,
Agamidae). Journal of Herpetology, 12 (4): 455-460.
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): 193-199.
Ota H. 1989. A new species of Japalura (Agamidae: Lacertilia: Reptilia)
from Taiwan. Copeia, 1989 (3): 569-576.
Ota H, Chen SL, Shang G. 1998. Japalura luei: A new agamid lizard from
Taiwan (Reptilia: Squamata). Copeia, 1998 (3): 649-656.
Pan XF, Zhou W, Zhou YW, Wu F, Zhang Q. 2002. Amphibian and reptile in
Zhongdian area of northwest Yunnan. Sichuan Journal of Zoology, 21 (2):
88-91. (in Chinese)
Pope CH. 1935. The Reptiles of China: Turtles, Crocodilians, Snakes,
Lizards. Natural History of Central Asia. Vol. X. New York: American Mu-
seum of Natural History, 466-467.
Qi Y, Wan HF, Gu HJ, Wang YZ. 2011. Do displays and badges function in
establishing the social structure of male toad-headed lizards, Phrynocepha-
lus vlangalii. Journal of Ethology, 29 (2): 381-387.
Reuter HI, Nelson A, Jarvis A. 2007. An evaluation of void filling interpola-
tion methods for SRTM data. International Journal of Geographic Informa-
tion Science, 21(9): 983-1008.
Sabaj Perez MH. 2015. Standard symbolic codes for institutional resource
collections in herpetology and ichthyology: an online reference. Version 5.0
Washington, DC. Available from: http: //www.asih.org/ (8 September 2015).
Schleich HH, Kästle W. 2002. Amphibians and Reptiles of Nepal. Ruggell:
Koeltz Scientific Books, 1-1201.
Schulte JA, Macey JR, Pethiyagoda R, Larson A. 2002. Rostral horn
evolution among agamid lizards of the genus Ceratophora endemic to Sri
Lanka. Molecular Phylogenetics and Evolution, 22 (1): 111-117.
Stuart-Fox DM, Ord TJ. 2004. Sexual selection, natural selection and the
evolution of dimorphic coloration and ornamentation in agamid lizards.
Proceedings of the Royal Society of London, Series B: Biological Sciences,
271 (1554): 2249-2255.
Townsend T, Mulcahy DG, Noonan BP, Sites JW Jr Kuczynski CA, Wiens JJ,
Reeder TW. 2011. Phylogeny of iguanian lizards inferred from 29 nuclear loci
and a comparison of concatenated and species-tree approach for an ancient,
rapid radiation. Molecular Phylogenetics and Evolution, 61(2): 363-380.
Wang K, Jiang K, Pan G, Hou M, Siler CD, Che J. 2015. A new species of
Japalura (Squamata: Sauria: Agamidae) from Eastern Tibet, PR China.
Asian Hepetological Research, 6 (3): 159-168.
Wei SY, Lin JY. 1981. Behavioral study of Japalura swinhonis formosensis
(Sauria: Agamidae). Tunghai Journal of Biology, 22: 33-48.
Xie Y, Mackinnon J, Li D. 2004. Study on biogeographic divisions of China.
Biodiversity and Conservation, 13(7): 1391-1417.
Xu J, Zhang WW. 2011. A field guide to the wildlife of Meili Snow Mountain
National Park. Beijing: Encyclopedia of China Publishing House, 1-400. (In
Chinese)
Yang DT, Rao DQ. 2008. Amphibia and Reptilia of Yunnan. Kunming:
Yunnan Publishing Group Corporation, 1-411. (in Chinese)
Zhao EM, Jiang YM. 1977. A survey of reptiles in Xizang Autonomous
Region, with faunal analysis and descriptions of new forms. Acta Zoological
Sinica, 23 (1): 64-71. (in Chinese)
Zhao EM, Zhao KT, Zhou KY. 1999. Fauna Sinica, Reptilia, Vol. 2: Squa-
mata, Lacertilia. Beijing: Science Press, 1-394. (in Chinese)
APPENDIX
The following specimens were examined:
J. batangensis (n=16): CIB 2227, 2233, 2243, 1902-1908, KIZ 84011,
801081, Batang, Sichuan, P.R. China; KIZ 019314, KIZ 09404, 019311,
019312, Markam, Tibet, China.
J. dymondi (n=7): CIB 87234, 1869, Panzhihua, Sichuan, P.R. China; KIZ
95I1001, 1002, 1016, 1018, 1022, Dayao, Yunnan, China.
J. grahami (n=1): USMN 65500 (holotype), Yibin, Sichuan, China.
J. micangshanensis (n=9): CIB 86351, 86348, Xianyang, Shaanxi, P.R.
China; CIB 86360, 86361, 86356, 86357, Luonan, Shaanxi, China; CIB
2572, 2578, 2582, Wenxian, Gansu, China.
J. flaviceps (n=13): CIB 2234, 2332, 2333, 2341, 2354, 2355, 2549, 2554,
2556, 2561, 2567; KIZ 05181, 05182; Luding, Sichuan, China.
J. splendida (n=6): USNM 35522 (holotype), Yichang, Hubei, PR China;
CIB 2588, 2591, 2596, 72468, 72469, Chongqing, China.
J. varcoae (n=3): CIB 2651, 2650, KIZ 85II0006, Kunming, Yunnan, China.
J. vela (n=11): KIZ 013801 (holotype), KIZ 013802, 013813, 013800,
013805–013811 (paratopotypes), Jerkalo, Tibet, China.
J. yunnanensis (n=8): CIB 2684, 2686, 2687, 2689, KIZ 82081, Longling,
Yunnan, PR China; KIZ 74II0240, 0248, 79I469, Tengchong, Yunnan,
China.
J. zhaoermii (n=12): CIB 86432, 86435, 85721, 85722, 86433, 86434,
86436, Wenchuan, Sichuan, PR China; CIB 2232, 2244, 2240, KIZ
84032, 85030, Lixian, Sichuan, China.