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Asian Herpetological Research 2015, 6(3): 159–168
DOI: 10.16373/j.cnki.ahr.140042
1. Introduction
The agamid lizards of the genus Japalura Gray, 1853 are
widely distributed in Asia from northwest India eastward
to Japan. Currently, the genus contains 28 recognized
species (Manthey et al., 2012; Uetz and Hošek, 2014),
with 14 of them recorded in mainland China: J.
andersoniana Annandale, 1905; J. batangensis Li et al.,
2001; J. brevicauda Manthey et al., 2012; J. dymondi
(Boulenger, 1906); J. fasciata Mertens, 1926; J. aviceps
Barbour and Dunn, 1919; J. grahami (Stejneger, 1924);
J. kumaonensis (Annandale, 1907); J. micangshanensis
Song, 1987; J. splendida Barbour and Dunn, 1919; J.
varcoae (Boulenger, 1918), J. yulongensis Manthey et al.,
2012; J. yunnanensis Anderson, 1878; and J. zhaoermii
Gao and Hou, 2002.
Among these species found in Mainland China, J.
flaviceps was reported from Chamdo (=Changdu) and
Markam (=Mangkang) of Chamdo Prefecture in eastern
Tibet (Hu et al., 1987; Li et al., 2010; Pope, 1935; Zhao
and Jiang, 1977). However, because later examinations of
historically collected specimens of Japalura restricted the
distribution of J. aviceps to the Dadu River Valley only
(Gao and Peng, 2005; Manthey et al., 2012), and since
Manthey et al. (2012) stated that the previously identied
specimen of J. flaviceps from the upper Lancang
(=Mekong) River at Jerkalo (=Yanjing) of Tibet may
represent an undescribed species, the taxonomic statuses
of Tibetan populations of J. aviceps are questionable.
A New Species of Japalura (Squamata: Sauria: Agamidae) from
Upper Lancang (Mekong) Valley of Eastern Tibet, China
Kai WANG
1, 2
, Ke JIANG
1
, Gang PAN
3
, Mian HOU
4
, Cameron D. SILER
2
and Jing CHE
1*
Abstract A new species of the agamid genus Japalura is described based on 15 specimens from the upper Lancang
(Mekong) Valley of eastern Tibet, PR China. Populations of the new species, Japalura vela sp. nov., were previously
recognized as J. aviceps. The new species is morphologically most similar to J. batangensis, J. micangshanensis, J.
variegata, and J. zhaoermii, but is distinguished from the four species and all remaining congeners by the following
combination of morphological characters: 1) small adult size (SVL 56–69 mm in males, 59–66 mm in females); 2)
ratio of tail TAL/SVL 1.85–2.06; 3) ratio of hind limb HLL/SVL 0.72–0.81; 4) T4S 24 or 25; 5) concealed tympanum;
6) transverse gular fold present; 7) gular pouch present; 8) axillary fold present; 9) a pronounced, continuous, sail-like
vertebral crest along length of body in males; 10) ground dorsal coloration black in males; 11) distinct gray transverse
streaks on dorsal surface of head; 12) black radiated streaks around eyes; 13) distinct, black vermiculate stripes on
ventral surface of head in both sexes; 14) a strongly jagged dorsolateral stripe from neck to base of tail on each side of
vertebral crest in males; and 15) absence of gular spots in both sexes. General distribution patterns of the genus in the
Hengduan Mountains region are also discussed.
Keywords distribution, Hengduan Mountains, Japalura, J. aviceps, species complex
*
Corresponding author: Prof. Jing CHE, from Kunming Institute of
Zoology, Chinese Academy of Sciences, Kunming, Yunnan, China,
with her research focusing on systematics, molecular evolution, and
biogeography of Asian amphibians and reptiles.
E-mail: chej@mail.kiz.ac.cn
Received: 28 July 2014 Accepted: 22 May 2015
ORIGINAL ARTICLE
1
State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of
Sciences, Kunming 650223, Yunnan, China
2
Sam Nobel Oklahoma Museum of Natural History and Department of Biology, University of Oklahoma, Norman
73072-7029, OK, USA
3
Institute of Plateau Ecology, Tibet Agriculture and Animal Husbandry College, Linzhi 860000, Tibet, China
4
Academy of Continuing Education, Sichuan Normal University, Chengdu 610068, Sichuan, China
Asian Herpetological Research160
Vol. 6
During a herpetofaunal survey of Tibet in 2013, we
collected 15 specimens of lizards identied to the genus
Japalura from the upper Lancang Valley at Quzika (near
Jerkalo) of the Chamdo Prefecture. In additional to a
number of morphometric differences, these specimens can
be distinguished easily from J. aviceps by the presence
of distinct radial streaks of pigmentation around the
eyes. Following robust morphological comparisons, this
population could not be assigned to any known congener.
Therefore, we describe this lineage as a new species
of Japalura and comment on the unique geographic
distributions of members of the genus in the Hengduan
Mountain regions.
2. Materials and Methods
All together 15 specimens of the new species were
collected from the upper Lancang Valley at Quzika,
eastern Tibet of China, including 10 adult males, two
adult females, and three juveniles. 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
eldwork. With the exception of a single male possessing
an incomplete tail (KIZ013798), the remaining 11
adult specimens were designated as the type series. All
specimens were deposited in the museum of the Kunming
Institute of Zoology, Chinese Academy of Sciences.
We measured morphological characters with a slide
caliper to the nearest 0.1 mm, except for the snout-vent
length (SVL) and tail length (TAL), which were measured
by a ruler to the nearest 1 mm, following the denitions of
Ota (1989). The following abbreviations of morphological
characters were used: snout–vent length (SVL), tail
length (TAL), head length (HL), head width (HW),
snout–eye length (SEL), orbital radius (OR), interorbital
distance (IOD), fore-limb length (FLL), hind limb length
(HLL), supralabial number (SL), infralabial number (IL),
subdigital lamellae number beneath Toe IV (T4S), and
mid-dorsal crest scale number, (MD). In addition to these
morphological characters, the following morphometric
characters were also examined: nasal–first supralabial
scale count (NSL), scale number between the nasal and
the first supralabial (NSL); supraciliary count (SCL),
elongated and overlapping scale number in a continuous
series from the posteriordorsal boundary of the nasal to
the point in line with the posterior margin of the orbit;
and supralabial–orbit scale row count (SOR), number of
horizontal scale rows between the sixth supralabial and
granular scales of the orbit right below the eye; length of
Toe IV (T4L), measured from the base of Toe IV to its
tip, excluding the claw; trunk length (TRL), measured
from the posterior insertion of fore-limbs to the anterior
insertion of hind limb. Values of paired characters from
both sides of the body are given in left/right order.
Voucher specimens of mainland species of Japalura
were examined (Appendix 1), including J. batangensis, J.
dymondi, J. aviceps, J. grahami, J. micangshanensis, J.
splendida, J. varcoae, J. yunnanensis, and J. zhaoermii.
In addition, data of the following ten species were also
obtained from Manthey et al. (2012): J. andersoniana,
J. brevicauda, J. chapaensis, J. fasciata, J. hamptoni, J.
otai, J. planidorsata, J. sagittifera, J. variegata, and J.
yulongensis. Morphometric data of the following nine
species were obtained from additional studies: J. brevipes
(Ota, 1989b), J. batangensis (Li et al., 2001; Wu et al.,
2004), J. kumaonensis
(Schleich and Kästle, 2002), J.
luei (Ota et al., 1998), J. makii (Ota, 1989a), J. swinhonis
(Ota, 1989b), J. tricarinata (Schleich and Kästle, 2002),
and J. zhaoermii (Gao and Hou, 2002). Coloration data of
live individuals of known Japalura species was obtained
from primary literatures listed above or published books
(Manthey, 2010; Schleich and Kästle, 2002; Yang and
Rao, 2008; Zhao et al., 1999). The museum abbreviations
referenced in this study include: Smithsonian National
Museum of Natural History (NMNH), Chengdu Institute
of Biology, Chinese Academy of Sciences (CIB),
Kunming Institute of Zoology, Chinese Academy of
Sciences (KIZ), and China West Normal University
(CWNU).
3. Results
Japalura vela sp. nov. Wang, Jiang, Che (Figures 1–6)
Synonymies
Japalura yunnanensis Vogt, 1924: 338
Japalura aviceps Hu et al., 1987: 112
Japalura aviceps Pope, 1935: 467
Japalura aviceps Zhao and Jiang, 1977: 293 –298
Japalura aviceps Zhao et al., 1999: 111–115
Japalura aviceps Li et al., 2010: 115
Japalura sp. A Manthey et al., 2012
Holotype KIZ013801 (Figures 1–4, 6), adult male from
Quzika of Markam, eastern Tibet, PR China (29°5' N,
98°36' E), at elevation of 2370 m, collected by Ke JIANG
and Kai WANG on May 23
rd
, 2013.
Paratypes Two adult females (KIZ013802 and
KIZ013813) and eight adult males (KIZ013800 and
KIZ013805–013811) all share the same data as the
holotype, collected by Ke JIANG, Kai WANG, and
Kai WANG et al. New Species of Japalura from TibetNo. 3 161
Duan YOU.
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) presence of enlarged crest
scales, 3) presence of gular sac, 4) presence of lateral
fold of skin in axilla–groin region, 5) supraciliary scales
greatly imbricate; 6) head relatively long, at; 7) tail long,
slender; 8) tail cylindrical in shape; and 9) absence of
precloacal or femoral pores.
The new species differs from all known congeners by
the following combination of characters: 1) small adult
size (SVL 56–69 mm in males, 59–66 mm in females);
2) moderate tail length (TAL/SVL 1.92–2.06 in males,
1.85–1.86 in females); 3) moderate hind limb length
(HLL/SVL 0.72–0.81); 4) T4S 24 or 25; 5) tympanum
concealed; 6) transverse gular fold present; 7) gular
pouch present; 8) axillary folds present; 9) males with
pronounced, continuous, sail-like vertebral crest along
entire length of body from posterior margin of head to
base of tail; 10) ground body coloration black in males,
medium to dark brown in females; 11) white coloration
on ventral surface of body in males; 12) presence of white
transverse streaks on dorsal head; 13) presence of black
radiated streaks around eyes; 14) ventral surface of head
with prominent black vermiculate stripes; 15) gular spots
absent; 16) presence of distinct, jagged, yellowish-white
dorsolateral stripes in males; and 17) presence of faint,
reddish, dorsolateral lines restricted to anterior half of
axilla–groin region in females.
Description of holotype Adult male, SVL 69 mm, TAL
136 mm, HL 20.8 mm, HW 14.5 mm, FLL 31.2 mm,
HLL 50.8 mm, T4L 13.2 mm, TRL 33.0 mm. Rostral
rectangular, four times longer than height, in contact with
nine scales including supralabials; nasal sub-rectangular,
separated from rostral by two small scales; supralabials
nine on both sides of head, weakly keeled, rst supralabial
separated from nasal by single small scale; scales in loreal
region irregularly arranged; small ciliaries around eyes,
separated from SL by three rows of scales; supraobital
ridge with single row of elongated, strongly keeled,
imbricate canthals and supraciliaries; scales posterior
to orbit greatly enlarged compared to loreals, strongly
keeled; tympanum concealed, covered with small scales.
Scales on dorsal surface of head heterogeneous in
size, strongly keeled, forming “Y”-shaped ridge from
postrostral to posterior edge of orbit. Mental pentagonal;
infralabials 10 on both sides of head, lacking keels;
scales on ventral surface of head homogeneous in size,
keel protuberance increasing posteriorly towards throat;
transverse gular fold present; gular pouch present; scales
covered by gular fold smaller than those on ventral
surface of head; shoulder fold posterior to gular fold on
each side of body.
Middorsal scales serrated, consisting of 46 relatively
large scales; in life, males with pronounced, sail-
like, continuous vertebral crest along entire length
of body from posterior margin of head to base of
tail; in preservation, crest reduced in protuberance;
dorsolateral ridge composed of large, strongly keeled
Figure 1 Dorsolateral close-ups, dorsolateral overviews, and ventral
overviews of Japalura vela sp. nov.: The male holotype KIZ013801
(A, B, and C) and the female paratype KIZ013802 (D, E, and F) in
life. Images not to scale. Photos by Kai WANG.
Figure 2 Dorsal (left) and ventral (right) views of holotype
(KIZ013801) of Japalura vela sp. nov. in preservative. Photo by
Kai WANG.
Asian Herpetological Research162
Vol. 6
scales continuous along length of body, centered along
dorsolateral stripes on each side; dorsal ground scales
small, slightly keeled, randomly distributed; lateral
ground scales similar to dorsal ground scales in size,
shape, keel protuberance, distribution; large, distinctively
keeled scales scattered among ground scales on dorsal
Figure 3 Dorsal head views of males of Japalura vela sp. nov. holotype KIZ013801 (A) and J. batangensis CIB1902 (B), J. aviceps
CIB2333 (C), J. zhaoermii CIB85722 (D), and J. micangshanensis CIB2572 (E) in preservative. Images not to scale. Photos by Ke JIANG.
Figure 4 Ventral head views of males of Japalura vela sp. nov. holotype KIZ013801 (A) and paratype KIZ013805 (B), J. batangensis
CIB1902 (C), J. aviceps CIB2333 (D), J. zhaoermii CIB85722 (E), and J. micangshanensis CIB2572 (F) in preservative. Images not to
scale. Photos by Ke JIANG and Kai WANG.
Figure 5 Ventral head views of females of Japalura vela sp. nov. paratype KIZ013802 (A), J. batangensis CIB1908 (B), J. flaviceps
CIB2234 (C), J. zhaoermii CIB86432 (D), and J. micangshanensis CIB86356 (E) in preservative. Photos by Ke JIANG.
Figure 6 Dorsal views of males of Japalura vela sp. nov. holotype KIZ013801 (A), J. aviceps KIZ05181 (B), and J. batangensis CIB2233 (C)
in preservative. Photos by Ke JIANG and Kai WANG.
Kai WANG et al. New Species of Japalura from TibetNo. 3 163
and lateral surfaces of body.
Scales on ventral surface homogeneous in size, greatly
keeled compared to dorsal ones. Limb scales more or
less homogeneous in size, keeled, more pronounced on
ventral surface; Toe IV subdigital lamellae 24. Tail scales
homogeneous in size, strongly keeled, keels align in
longitudinal rows along entire length of tail.
Coloration of holotype The dorsal surface of the head
is black with three distinct, narrow, ash gray transverse
streaks, one in a “m”-shape between the nares, and two
in a “X”-shape between the eyes. The lateral surface of
the head is white with nine or ten black streaks radiating
around each eye. Four of these streaks are observed below
the eyes on both sides of the head, and extend to the
supralabial scales, with the posteriormost streak being the
broadest. The infralabial scales have short black bars that
are aligned with streaks on the supralabials. Pigmentation
on the chin and throat is white with distinct black
vermiculate stripes angled medially towards the midline
of the body. Dorsal and lateral surfaces of the body are
black, but gradually fade to brownish black from the base
of the tail to its tip, as well as from the proximal to the
distal end of the limbs. Prior to capture, the individual
possessed a strongly jagged yellow dorsolateral stripe
on each side of the vertebral crest along the length of
the body. Following capture, the coloration of the two
stripes changed to yellowish brown. Seven narrow yellow
bands are present across the dorsal surface of the body
from the neck to the groin. Posterior bands 3–7 extend
across the dorsolateral stripes on both sides of the body.
Lateral surface of the body in the axilla–groin region
possesses irregularly distributed, small to moderate-
sized, yellow spots. The larger lateral yellow spots nearly
form two longitudinal rows. The dorsal surface of the tail
is brownish black with 16 light-gray, transverse bands
evenly distributed along the length of the tail. Coloration
of the dorsal surfaces of limbs consists of a brownish
black ground color with white transverse bands running
along their length. The ventral surfaces of the body and
limbs are uniformly white, except the palms and soles,
which are yellowish-white (Figure 1).
Although the coloration of the holotype in preservation
closely matches its coloration in life, the following
differences were observed: 1) dorsolateral stripes are dark
gray, 2) transverse bands on the dorsal surface of the body
and spots on lateral surface of the body are white, and 3)
dorsal surface of the tail is dark gray (Figure 2).
Variation Variation in morphometric and meristic
characters is summarized in Table 1. Additionally,
the following coloration differences exist in the male
paratypes: 1) three male paratypes (KIZ013805, 013806,
and 013809) have signicantly faded bluish coloration on
the gular pouches, but nevertheless the faded color does
not form clear gular spots (Figure 4); 2) several dorsal
transverse bands do not extend completely across the
dorsal midline in three males (KIZ013808–10); and 3)
the vertebral skin fold of one male (KIZ 013800) is less
developed than the holotype.
Sexual dimorphism is observed for a number of
characters. Females have more vertically compressed
bodies (vs. less compressed), shorter tails (ratio of
TAL/SVL 1.85–1.86 vs.
≥
1.92), highly reduced vertebral
crests (vs. pronounced and sail-like), brown ground
coloration on the dorsal and lateral surfaces of the body
(vs. black), a greater number of keeled, paravertebral
scale rows on dorsal and lateral surfaces of body (four vs.
two), and a faint, reddish dorsolateral lines restricted to
anterior half of axilla–groin region (vs. a distinct yellow
stripes along the entire length of the body). Additionally,
females possess chevron-shaped brown pigmentation
blotches along the dorsal midline of the body (vs. absent),
and white, dorsal, transverse bands that extend across
the lateral portions of the body (vs. extension just past
the dorsolateral stripes; Figure 1). Overall, the juveniles
(KIZ013799, 013804, 013812) resemble the adult
females, but they differ by having granular scales on the
dorsal and lateral surfaces of the head, broader, white
transverse bands on the dorsal surface of body, and by the
absence of unclear reddish dorsolateral lines.
Comparisons Japalura vela sp. nov. was previously
recognized as J. aviceps (Hu et al., 1987; Li et al., 2010;
Zhao and Jiang, 1977), but it can be distinguished easily
from the latter by having a greater number of T4S (24 or
25 vs. 21–23), a smaller adult body size (maximum SVL
up to 69mm vs. to 83mm), a relatively shorter snout (ratio
of SEL/HL 0.33–0.38 vs. 0.39–0.44), a pronounced, sail-
like, and continuous vertebral crest in males (vs. weak or
discontinuous with a clear break in the skin fold between
the nuchal and dorsal regions of the body), a black dorsal
background coloration in males (vs. brownish gray), black
(in males) or brown (in females) rectangular patches on
the dorsal surface of the body along the midline (vs. a
lateral series of dark brown rhombs with yellow centers),
as well as by the presence of multiple distinct transverse
streaks on the dorsal surface of the head (vs. absent,
signicantly faded, or in low numbers), and the presence
of distinct radiated streaks around the eyes (vs. absent).
The new species is morphologically most similar to
J. batangensis, J. micangshanensis, J. zhaoermii, and J.
variegata in external morphology, with all five species
Asian Herpetological Research164
Vol. 6
having transverse streaks on the dorsal surface of the head,
distinct radiated streaks around the eyes, pronounced
vertebral crest in males, and dorsolateral stripes that run
parallel to the dorsal midline in males. However, males
of the new species can be distinguished from males of
the latter four species by having a pronounced, sail-like,
continuous vertebral crest running along the entire length
of the body, with no distinct break between the nuchal and
dorsal sections (vs. discontinuous or having a clear break
between the two sections). Additionally, the new species
differs from J. batangensis by having a greater number
of T4S (24 or 25 vs. 18–22), a tendency towards longer
hind limbs (ratio of HLL/SVL 0.72–0.81 vs. 0.65–0.75),
a white coloration on ventral surface of body in males
(vs. bright yellow), as well as by the presence of unclear
reddish lines dorsolaterally in females (vs. absent or
presence of distinct dorsolateral stripes), and the absence
of a distinct gular spot (vs. always present in males, often
in females, greenish blue); from J. micangshanensis by
having a black dorsal background coloration in males (vs.
brownish), as well as by the presence of a gular fold (vs.
absent), the presence of distinct black vermiculate stripes
on the ventral surface of the head (vs. absent), and by the
presence of unclear reddish lines dorsolaterally in females
in life (vs. absent); from J. zhaoermii by having a smaller
adult body sizes (maximum SVL up to 69 mm vs. to 85
mm), as well as by the absence of a gular spot (vs. present
in males, yellowish green), and the presence of distinct,
narrow, black vermiculate stripes on ventral surface of
heads in males (vs. indistinct, broad stripes that fade
signicantly towards the center of the gular pouch); and
from J. variegata by having fewer MD (41–51 vs. 54–67),
a black dorsal background coloration (vs. olive green),
as well as by the absence of black reticulated patterns on
the lateral surface of the body (vs. present), and by the
absence of a gular spot (vs. present, violet).
Japalura vela sp. nov. can also be distinguished from
the congeners that distribute along the same river or in
close proximity. The new species can be distinguished
from J. yunnanensis, which occupies the lower reaches
of Lancang River, by having a shorter tail (ratio of TAL/
SVL
≤
2.06 vs.
≥
2.25), one or two scales between nasal
and the first SL (vs. in direct contact), a black ground
coloration of the dorsal surface of the body in males
(vs. brown or olive green), as well as by the absence of
a distinct gular spot in males (vs. present in males, light
yellow), and the presence of prominent black vermiculate
stripes on the ventral surface of the head (vs. absent). For
species from adjacent areas, the new species differs from
J. brevicauda by having a longer tail (ratio of TAL/SVL
Number Status Sex SVL TAL TAL/SVL HL HL/SVL HW HW/SVL SEL IOD FLL FLL/SVL HLL HLL/SVL T4L TRL SL IL NSL MD T4S SOR
SCL
KIZ013801 holotype male 69 136 1.97 20.8 0.3 14.5 0.21 7.4 9.5 31.2 0.45 50.8 0.74 13.2 33.0 9/9 10/10 1/1 46 25/25 3/3
7/7
KIZ013809 paratype male 60 115 1.92 18 0.3 12.8 0.21 6.4 8.7 28.4 0.47 44.9 0.75 11.3 25.3 8/8 10/11 1/1 42 24/25 3/3
7/7
KIZ013808 paratype male 65 134 2.06 19.5 0.3 14 0.22 6.9 9.6 31.5 0.49 50.3 0.77 12.3 31.3 8/8 9/10 2/2 47 24/25 3/3
7/7
KIZ013800 paratype male 62 122 1.97 18.7 0.3 13.5 0.22 6.8 9.3 29.9 0.48 50.1 0.81 12.0 29.4 8/8 11/11 2/1 41 24/24 3/3
7/7
KIZ013805 paratype male 60 123 2.05 18.8 0.31 13 0.22 7 9.2 28.6 0.48 43.9 0.73 11.9 29.0 9/10 11/10 1/2 44 25/25 4/4
7/7
KIZ013807 paratype male 56 110 1.96 17.3 0.31 12.2 0.22 6.5 8.4 28.0 0.50 44.0 0.79 11.3 26.0 9/9 10/10 1/1 44 24/24 3/3
7/7
KIZ013806 paratype male 65 129 1.98 20 0.31 13.6 0.21 7.4 9.6 31.5 0.48 46.6 0.72 12.6 29.7 9/10 10/10 1/1 44 25/24 3/3
7/7
KIZ013811 paratype male 64 127 1.98 19.9 0.31 14.1 0.22 6.9 9.2 29.3 0.46 45.8 0.72 13.1 28.6 7/8 9/9 1/2 44 24/25 3/3
7/7
KIZ013810 paratype male 57 112 1.96 19.2 0.34 13.9 0.24 6.4 9 25.5 0.45 44.7 0.78 11.9 26.3 9/9 11/11 2/2 45 24/24 4/4
7/7
Range 56–69 110–136 1.92–2.06 17.3–20.8 0.30–0.34 12.2–14.5 0.21–0.24 6.4–7.4 8.4–9.6 25.5–31.5 0.45–0.50 43.9–50.8 0.72–0.81 11.3–13.2 25.3–33.0 7–10 9–11 1–2 41–47 24–25 3 or 4
7
Average 62 123 1.98 19.1 0.31 13.5 0.22 6.9 9.2 29.2 0.47 46.9 0.76 12.2 28.8 8.61 10.17 1.39 44.1 24.4 3.22
7
KIZ013802 paratype female 66 123 1.86 19.5 0.3 14 0.21 7.4 9.1 32.0 0.48 47.9 0.73 13.9 32.7 9/9 11/10 2/2 51 24/24 3/3
8/8
KIZ013813 paratype female 59 109 1.85 17.3 0.29 12.4 0.21 6.2 8.8 28.0 0.47 43.1 0.73 12.1 30.2 9/9 10/11 2/1 47 25/25 3/3
7/8
Average 63 116 1.86 18.4 0.3 13.2 0.21 6.8 9 30.0 0.48 45.5 0.73 13.0 31.5 9 10.5 1.75 49 24.5 3
7.33
Table 1 Morphometric characters and meristic data of the type series of Japalura vela sp. nov.
Measurements of SVL, TAL, HL, HW, T4L, TRL, FLL, and HLL are in the unit of millimeter (mm), all meristic data are given in decimals, and paired meristic characters are all given in left/right
order. For average calculation of paired measurements, each one of the paired measurements is treated independently. Full terms of abbreviations are shown in the method section.
Kai WANG et al. New Species of Japalura from TibetNo. 3 165
≥
1.85 vs.
≤
1.45), longer fore-limbs (ratio of FLL/SVL
≥
0.45 vs.
≤
0.40), longer hind limbs (ratio of HLL/SVL
≥
0.72 vs.
≤
0.64), greater number of T4S (24 or 25 vs.
16–20), as well as by the presence of a distinct gular
pouch (vs. not visible); and from J. yulongensis by having
a shorter tail (ratio of TAL/SVL ratio < 2.06 vs. >2.23), as
well as by the absence of a distinct gular spot in preserved
male specimens (vs. present, dark in preservation).
The new species J. vela sp. nov. can be easily
distinguished from the remaining mainland congeners
by having distinct ornamentations and obvious
morphological characteristics without detailed
comparisons of pholidosis. The new species differs from
J. andersoniana by having shorter hind limbs (ratio of
HLL/SVL 0.72–0.81 vs. 0.88–1.10); from J. fasciata by
the absence of pale-blue, hourglass-shaped pattern on mid
body (vs. present); from J. grahami by having a greater
number of MD (41–45 vs. 10); from J. otai Mahony,
2009, J. sagittifera Smith, 1940, and J. planidorsata
Jerdon, 1870 by having a un-depressed body shape (vs.
vertically depressed), as well as by the presence of two
distinct, broad dorsolateral stripes in males (vs. absence or
distinctively narrow), and the absence of a light vertebral
strip on the dorsal surface of the body (vs. present); from
J. dasi, J. dymondi, J. kumaonensis, J. tricarinata (Blyth,
1854), and J. varcoae by having a concealed tympanum
(vs. exposed); from J. major (Jerdon, 1870) by having a
concealed tympanum (vs. mostly exposed), a black ground
coloration with white spots on the lateral surface of the
body in males (vs. olive ground color, reticulated with
black on the lateral sides), a uniform white coloration on
ventral surface of the body (vs. speckled with black), as
well as by the absence of white lip-stripes below the eyes
(vs. present); from J. splendida by having a smaller adult
body size (maximum SVL < 69 mm vs. <92 mm), jagged
dorsolateral stripes in males (vs. with smooth edges),
a black (males) or brown (females) ground coloration
of the dorsal surface of the body (vs. gray or green), as
well as by the presence of pronounced, continuous, sail-
like vertebral crest in males (vs. feebly developed and
discontinuous); from J. hamptoni Smith, 1935 by having
parallel, strongly jagged dorsolateral stripes in males (vs.
diagonally away from the dorsal midline and with smooth
edges); and from J. chapaensis Bourret, 1937 by having
fewer T4S (24 to 25 vs. 28), a light-pink coloration of the
oval cavity in life (vs. yellow), as well as by the absence
of distinct gular spot (vs. present in males, yellow), and
the absence of reticulated black pigmentation on the
lateral surface of the body (vs. present).
Additionally, the new species differs from all island
species (J. breviceps Gressit, 1936; J. luei Ota, Chen
and Shang, 1998; J. makii Ota, 1989; J. polygonata
polygonata (Hallowell, 1860); J. polygonata donan Ota,
2003; J. polygonata ishigakiensis Van Denburgh, 1912;
J. polygonata xanthostoma Ota, 1991; and J. swinhonis
Günther, 1864) by having a transverse gular fold (vs.
absent), a black ground coloration of the dorsal surface
of the body in males (vs. brown or green), the parallel
dorsolateral stripes (vs. absent or diagonally away from
dorsal midline), and a terrestrial lifestyle (vs. arboreal),
as well as by the absence of a distinct gular spot in males
(vs. present).
Distribution and Ecology The new species is currently
known only from the type locality (Figures 7–8), but
it may be found in valleys of adjacent reaches along
Lancang Rivers. As a terrestrial species, individuals were
observed commonly in rocky areas or steppe-shrub habitat
along the arid river valley (Figure 7). Adult males usually
basked on high rocks, while adult females and juveniles
stayed lower in the rock piles, suggesting possible niche
Figure 7 The microhabitat (A) and macrohabitat (B) of Japalura
vela sp. nov. at the type locality, Quzika, Tibet. Photos by Duan
YOU.
Asian Herpetological Research166
Vol. 6
partitioning among different age-groups and between
different sexes. Males are territorial, in which the territory
holder will perform vertical head-nodding movements
and display gular pouch toward the invader, and physical
contacts (biting and chasing) will happen if the invader
refuses to leave. No territorial behaviors were seen among
females or juveniles. Possible predations may come from
snakes (Chinese Beauty Snake, Orthriophis taeniurus,
KIZ013803, was collected from the same locality) and
large birds (Corvus sp., also commonly observed at
this locality).
Etymology The Latin word vela means “sail”, which
describes the shape of the pronounced and continuous
vertebral crest as the diagnostic morphology of the males
of the new species. Hence according to the Latin name,
we suggest Sail Moutain Lizards or Sail Japalura as its
English common name, and Fan Bei Pan Xi (
帆背攀蜥
)
as its Chinese common name.
4. Discussion
Previous studies have suggested that Japalura lizards
show a strong tendency to distribute along river valleys in
the Hengduan Mountains, where each species is endemic
to specific reaches of the river (Manthey et al., 2012;
Yang and Rao, 1992). For example, along the Yangtze
River, J. batangensis is endemic to the very upper reaches
of Jinsha River (Li et al., 2001; Wu et al., 2005); J.
yulongensis and J. brevicauda have been found in slightly
lower reaches in northwestern Yunnan only (Manthey et
al., 2012); J. dymondi has been reported in further lower
reaches of north central Yunnan and southern Sichuan
(Yang and Rao, 2008; Zhao et al., 1999); and J. splendida
has been found in the rest of the middle reaches of
Yangtze River (Manthey et al., 2012; Zhao et al., 1999).
For the Lancang River, J. yunnanensis was reported to
occur in the lower reaches in southwestern Yunnan (Jiang,
1992; Yang and Rao, 2008; Zhao et al., 1999), and J.
vela appears to be distributed along the upper reaches of
the Lancang River in Tibet. This continuous distribution
pattern of species in the genus along the Lancang River
is consistent with the river-based distribution pattern of
other mainland congeners in East Asia. Given the large
number of un-surveyed river habitats and additional
unexamined records J. cf. flaviceps in Yunnan Province
(Pan et al., 2002; Yang and Rao, 2008), there may still be
undiscovered species diversity in the valleys of the upper
Lancang River Basin and adjacent areas.
The rise of the Hengduan Mountains have
been shown to have created geographic barriers to
formerly continuous populations of species within the
herpetofauna, which resulted in population diversication
and, subsequently, speciation (Che et al., 2010; Huang et
al., 2009; Yang et al. 1987). For the genus Japalura, J.
vela and J. batangensis occur in close proximity (about
60 km between their native valleys), and yet the two
river valleys are separated by continuous mountains
reaching over 3 500 m in elevation (Figure 8). Given that
the known upper distribution limit of Japalura in the
area is 3 000 m in elevation only (Yang et al., 1987), we
believe these two species to be allopatrically distributed.
Much work remains to be done with this genus, including
Figure 8 Map showing the relative positions of the type locality of Japalura vela sp. nov. (A) and J. batangensis (B) in PR China.
Kai WANG et al. New Species of Japalura from TibetNo. 3 167
studies on sexually dimorphic traits, crest evolution,
and biogeographic patterns. Future phylogeographic
and behavioral studies are needed to gain a better
understanding of their unique evolutionary history.
Acknowledgements This work was supported by
the Ministry of Science and Technology of China
(2014FY210200, 2011FY120200), the National Natural
Science Foundation of China (31090250), and the
Animal Branch of the Germplasm Bank of Wild Species
of Chinese Academy of Sciences (the Large Research
Infrastructure Funding). We thank Mr. Duan You, who
assisted us in the fieldwork in Tibet, Dr. Gernot Vogel
(Society for Southeast Herpetology, Germany), who
gracefully provided suggestions on the Latin name of this
new species. We also thank Prof. Yuezhao Wang (CIB),
Prof. Yueying Chen (CIB), Mr. Ke Lu (CIB), Mr. Guiwu
He (KIZ), and Mr. Wynn Addison (NMNH) for their
help in examining the specimens, and Ms. Jingting Liu
(Washington State University) for her help in editing the
photographs.
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Appendix 1 Specimens examined.
Japalura batangensis (n=19): CWNU98012 (holotype), CWNU98004, 98010, 98014–18, CIB2227, 2233, 2243, 1902–
09, Batang, Sichuan, China.
J. dymondi (n=3): CIB87234, 1869, 1817, Panzhihua, Sichuan, China.
J. aviceps (n=10): CIB2332, 2341, 2354, 2355, 2333, 2234, 2561, 2556, Luding, Sichuan, China; KIZ05181–05182,
Luding, Sichuan, China.
J. grahami (n=1): USMN65500 (holotype), Yibin, Sichuan, China.
J. micangshanensis (n=9): CIB86351, 86348, Xianyang, Shanxi, China; CIB86360, 86361, 86356, 86357, Luonan,
Shanxi, China; CIB2572, 2578, 2582, Wenxian, Gansu, China.
J. splendida (n=6): USNM 35522 (holotype), Yichang, Hubei, China; CIB2588, 2591, 2596, Wushan, Chongqin, China;
CIB72468, 72469, Yunyang, Chongqin, China.
J. varcoae: (n=2): CIB2651, 2650, Kunming, Yunnan, China.
J. yunnanensis (n=4): CIB2686, 2687, 2689, 2684, Longling, Yunnan, China.
J. zhaoermii (n=8): CIB002690 (holotype), CIB86435, 86432, 85722, 85721, Wenchuan, Sichuan, China; CIB2232,
2244, 2240, Lixian, Sichuan, China.
