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wileyonlinelibrary.com/journal/jzs J Zool Syst Evol Res. 2020;67:222–263.© 2020 Wiley-VCH GmbH
Received: 3 April 2020
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Revised: 9 July 2020
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Accepted: 12 July 2020
DOI: 10.1111/jzs.12414
ORIGINAL ARTICLE
Systematic revision of mountain dragons (Reptilia: Agamidae:
Diploderma) in China, with descriptions of six new species and
discussion on their conservation
Kai Wang1,2 | Jinlong Ren3 | Jiawei Wu4 | Ke Jiang3 | Jieqiong Jin2 |
Shaobing Hou2 | Puyang Zheng3 | Feng Xie3 | Cameron D. Siler1 | Jing Che2
Camer on D. Siler and Che J ing shou ld be considere d as joint senior au thors.
Contri buting autho rs: Jinlong Ren (r enjl@cib.ac .cn), Jiawei Wu (wuji awei@s wild.c n), Ke Jiang (3465990 56@qq. com), Jieqiong Ji n (jinjq@ma il.kiz .ac.cn), Shao bing Hou (1603371747@
qq.com), P uyang Zheng (2374 48947@qq.com), Feng X ie (xief eng@cib.ac .cn).
1Sam Noble Oklah oma Museum of Natu ral
Histor y and De part ment of Biology,
University of Oklahom a, Norm an, OK , USA
2State Key Laboratory of Genetic Resources
and Evolution, Kunming Institute of Zoolog y,
Chinese Acade my of Scien ces, Kunming,
Yunnan, China
3Chengdu Instit ute of Biol ogy, Chinese
Academy of Sciences, Che ngdu, Si chuan,
China
4Swild Studio, Chengdu, Sic huan, China
Correspondence
Kai Wang and Cameron D. Siler, Sam Noble
Oklahoma Museum of Natur al History
and Department of Biology, University of
Oklahoma, Nor man, Oklahoma, USA .
Emails: kai.wang-2@ou.e du (KW ) and
camsiler@ou.edu (CDS)
Jing Che, State Key Laboratory of G enetic
Resources and Evolution, Ku nming Institu te
of Zoolog y, Chinese Academy of Sciences,
Kunming, Yunnan, China.
Email: chej@kiz.ac.cn (JC)
Funding information
STEP Program of China, Gr ant/Award
Number : 2019QZKK0501; National Key
Research and deve lopment Program
of China, G rant/Award Numbe r:
2017YFC0505202; Strategic Priority
Research Program of CAS, China , Grant/
Award Number: XDA19050303; The
Ministry of Ecol ogy and Environment
of China, G rant/Award Numbe r:
2019HJ20960 01006; National Science
Foundat ion of USA , Grant/Award Num ber:
DEB 1657648, IOS 1353683, G RFP
2017216966 and EAP SI 171400
Abstract
Cryptic diversity not only introduces confusion to taxonomic studies, but it also
poses major challenges to conservation and environmental legislation. One such
troubling group are the Mountain Dragons in the genus Diploderma in Southwest
China. Previous studies have suggested that the genus contains considerable cryp-
tic diversity, particularly in the D. flaviceps complex. Owing to taxonomic confusion,
micro-endemic lineages are still neglected by the Chinese wildlife protection laws,
despite their urgent conservation needs. Combining multivariate morphological and
multi-locus phylogenetic data, we provide the first integrative systematic revision of
the genus Diploderma. Specifically, we confirm that the six examined populations of
D. cf. flaviceps from the upper Jinsha and Yalong River Valleys in Sichuan and Yunnan
Provinces represent six cryptic, undescribed lineages, and we describe each of them
as a new species. With the updated taxonomy and distribution information, we dis-
cuss the taxonomy of the D. flaviceps complex in Southwest China, provide an up-
dated diagnostic key along with distributional ranges for all species of the genus, and
discuss some of the suspicious records of other congeners in China. Lastly, we evalu-
ate the IUCN status of each of the six new species and highlight the major challenges
for Diploderma conservation in China due to delayed environmental legislation and
misleading conservation assessments.
KEYWORDS
Draconinae, IUCN, Japalura sensu lato, taxonomy, Wildlife Protection Act of China
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1 | INTRODUCTION
The genus Diploderma Hallowell, 1861, commonly known as
Mountain Dragons, was resurrec ted recently from the paraphyletic
genus Japalura sensu lato Gray, 1853 (Wang, Che, Lin, et al., 2019).
Despite the recent split, Diploderma still comprises a major portion of
the diversity in the subfamily Draconinae, with a total of 25 species
recognized to date ( Wang, Che, et al., 2019; Wang, Jiang, Ren, et al.,
2019; Wang, Wu, Jiang, et al., 2019). Most congeners are found in the
Hengduan Mountain Region (HMR) in Mainland Southwest China and
Taiwan (Manthey, 2010; Wang, Che, et al., 2019), and the major por-
tion of the diversity (12 species) belongs to a single species complex,
the D. flaviceps (Barbour & Dunn, 1919) complex (Figure 1; Manthey,
Wolfgang, Hou, & Wang, 2012; Wang, Jiang, Ren, et al., 2019).
First described based on specimens from the upper Dadu River
Valley in central Sichuan Province, China, the type specimen (ho-
lotype MCZ R-12469) of D. flaviceps was deposited in the United
States. With no access to the t ype specimen and logistical obsta-
cles to accessing the type locality, early Chinese herpetologists
misinterpreted the diagnosis of D. flaviceps, particularly regarding
the coloration and ornamentation patterns of the head and body.
In fact, incorrect diagnostic characters were widely recognized for
years in early Chinese literature, including the supposed presence
of greenish body coloration, colorful gular spots, and distinct radial
stripes around the eyes (Hu et al., 1987; Zhao, Zhao, & Zhou, 1999).
As a result, nearly all Diploderma populations in Southwest and cen-
tral China were long considered as D. flaviceps, a species believed to
have the widest distribution among all Chinese congeners (Yang &
Rao, 2008; Yao & Gong, 2012; Zhao et al., 1999).
Recent re-examination of D. c f. flaviceps populations across China
have restricted the range of true D. flaviceps to the upper Dadu River
Valley only, and studies have shown that the majority of congeners
from the HMR are micro-endemic lineages restricted to only specific
sections of isolated river valleys (Manthey et al., 2012; Wang et al.,
2015, 2016; Wang, Jiang, Ren, et al., 2019). Therefore, populations
of previously identified D. cf. flaviceps out side of the upper Dadu
River Valley represent a large species complex of misidentified con-
geners. Since the early 2000s, multiple distinct evolutionar y lineages
have been discovered from the D. flaviceps com plex across the HMR ,
including D. batangense (Li, Deng, Wu, Wang, 2001), D. brevicaudum
(Manthey, Wolfgang, Gou, Wang, 2012), D. drukdaypo (Wang, Ren,
Jiang, Zou, Wu, Che, Siler, 2019), D. iadinum (Wang, Jiang, Zou, Yan,
Siler, Che, 2016), D. laeviventre (Wang, Jiang, Zou, Yan, Siler, Che,
2016), D. micangshanense (Song, 1987), D. vela (Wang, Jiang, Pan,
Hou, Siler, Che, 2015), D. yulongense (Manthey, Wolfgang, Hou,
Wang, 2012), and D. zhaoermii (Gao, Hou, 20 02; Figure 1). However,
FIGURE 1 Distributions of Diploderma species in East Asia. Shaded region represents the range of the D. flaviceps complex in China.
Numbered shapes represent type localities of Diploderma species: The rectangle represents true D. flaviceps (5), stars represent the new
species described here (6: D. panchi sp. nov.; 7: D. angustelinea sp. nov.; 8: D. panlong sp. nov.; 12: D. flavilabre sp. nov.; 14: D. aorun sp. nov.;
and 15: D. qilin sp. nov.), and circles represent type localities of the remaining recognized congeners (1: D. splendidum; 2: D. micangshanense;
3: D. zhaoermii; 4: D. grahami; 9: D. swild; 10: D. dymondi; 11: D. varcoae; 13: D. batangense; 16: D. brevicaudum; 17: D. yulongense; 18:
D. drukdaypo; 19: D. vela; 20: D. iadinum; 21: D. laeviventre; 22: D. slowinskii; 23: D. yunnanense; and 24: D. chapaense)
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WANG et Al .
despite the well-recognized suspicion of taxonomic identities of
D. cf. flaviceps and the recent efforts in taxonomic studies of the
D. flaviceps complex, the status of many questionable populations of
D. cf. flaviceps in the HMR still remains unexamined. This is particu-
larly true for populations from the valleys of the Jinsha River and its
main tributar y, the Yalong River, in Yunnan and Sichuan Provinces
(Wang, Ren, Jiang, et al., 2019).
In Yunnan Province, Yang and Rao (2008) reported on question-
able records of D. cf. flaviceps from the Ji ns ha Val le y in Deqi n Co unt y
and Shangri-La County (=Xianggelila). Based on their descriptions,
these Yunnan populations of D. cf. flaviceps possess distinct dark ra-
dial stripes around the eyes and colorful gular spots, which contra-
dict the revised diagnoses of true D. flaviceps (radial stripes around
the eyes absent or faint; gular spots absent; Manthey et al., 2012;
Wang et al., 2015; Wang et al., 2016; Wang, Ren, Jiang, et al., 2019).
As coloration and ornamentation patterns have proved reliable as
diagnostic characters for members of the genus Diploderma (Wang
et al., 2015, 2016, 2017; Wang, Jiang, Ren, et al., 2019; Wang, Ren,
Jiang, et al., 2019; Wang, Wu, Jiang, et al., 2019), these Yunnan pop-
ulations with such contradicting ornamentation patterns warrant
further investigation.
In Sichuan Province, Zhao (20 03) recorded D. c f. flaviceps from
Mianning and Muli Counties, without providing any vouchered in-
formation of referred specimens; however, both of the mentioned
co u ntie s are al o ng the Yalo ng Ri ver Val ley, wh i c h is is ola ted fro m th e
restricted range of the true D. flaviceps (i.e., Dadu River Valley) by
the continuous Hengduan Mountains. Because these populations
of D. cf. flaviceps in Sichuan Province are from localities that are
allopatric from the known range of true D. flaviceps, separated by
known biogeographic barriers (i.e., Hengduan Mountains; Lei, Qu,
Song, Alström, & Fjeldså, 2015; Yang, Yi, Pan, & Guo, 2012), they
may also represent overlooked cryptic diversity within the genus.
Furthermore, in addition to the abovementioned suspect popu-
lations of D. cf. flaviceps, many suitable valleys in Sichuan Province
have not yet been surveyed for Diploderma diversity, including the
upper reaches of the Jinsha and Yalong Rivers in Baiyu and Yajiang
Counties, respectively. Given the consistent distributional patterns
of Diploderma re str icte d to th e maj or ri ve r s in th e HMR (Wan g, Ji ang ,
Ren, et al., 2019), these un-surveyed valleys also may harbor addi-
tional undescribed diversity.
To decipher cryptic diversity within the D. flaviceps species com-
plex , an int eg rat iv e ap proac h mu st be taken . Howe ve r, du e to the lack
of specimens and genetic samples, not only were the majorit y of the
previous taxonomic studies on the genus Diploderma based on mor-
phological data only (Manthey et al., 2012; Rao, Vindum, Ma, Fu, &
Wilkinson, 2017; Wang, Jiang, Ren, et al., 2019; Wang et al., 2017), but
the few recent integrative taxonomic studies of the genus also have
limited taxa or gene coverage (Wang, Che, et al., 2019; Wang, Ren,
Jiang, et al., 2019; Wang, Wu, Jiang, et al., 2019). Some recognized
species still lack molecular data, and their phylogenetic relationships
with congeners remain unknown. Such paucity of comprehensive ge-
netic data and phylogenetic relationships among recognized species
hinders the discover y processes of cryptic diversity in this group.
Lastly, the current unresolved taxonomy poses major challenges
to the conser vation of the genus Diploderma in China. Cryptic, un-
described species and species re cently discovered are left out of en-
dangered species listings, and taxonomic confusion has resulted in
misleading conservation assessments (Wang, Jiang, Ren, et al., 2019;
Wang, Ren, Jiang, et al., 2019). With the rapidly expanding human
settlements in the HMR, this lag between species discovery and en-
vironmental legislation and conservation action is an urgent concern
that needs to be addressed.
During herpetological surveys between 2015 and 2019, we
collected specimens of the genus Diploderma across the HMR in
Southwest China (denoted in stars numbered 6, 7, 8, 12, 14, and 15
in Figure 1). Based on these collections, we t ake the first comprehen-
sive, integrative approach to investigating diversity within the D. fla-
viceps complex in Southwest China, using multi-locus phylogenetic
and morphological analyses. Our results indicate that members of the
D. flaviceps complex are paraphyletic, and the six examined popula-
tions identified previously as D. cf. flaviceps from Yunnan and Sichuan
Provinces represent undescribed, micro-endemic diversity within the
genus that are facing urgent conservation needs. Herein, we describe
six new species of Diploderma from the D. flaviceps complex, provide
an updated diagnostic key of the genus, and discuss the possible solu-
tions to the major conservation challenges that these Chinese endemic
species face due to delayed conservation assessments and legislation.
2 | MATERIALS AND METHODS
2.1 | Taxonomic sampling
Specimens of the genus Diploderma were collected in southwest-
ern China from 2015 to 2019, including 54 specimens from six
populations of D. cf. flaviceps: three from the upper Jinsha River
Valley and three from the Yalong River Valley (Figure 1; for de-
tailed population sampling see Appendix 1 and Table S1). In the
field, specimens were euthanized, 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. All newly collected specimens
were deposited at the Museum of Kunming Institute of Zoology,
Chinese Academy of Sciences (KIZ).
In addition to specimens collected, specimens in major museum
collections were also examined, which included 223 specimens of 22
recognized species of the genus Diploderma (Tabl e S1). To avoid pos sibl e
taxonomic confusion over cryptic diversity, morphological data were
taken from type specimens, topotypic specimens, or specimens with
confirmed taxonomic status whenever possible. Museum abbreviations
fol low Sabaj (2016), wh ich inc lu de d the fol lowing: Mus eu m of Ca lifornia
Academy of Sciences (CAS), San Francisco, CA, USA; Chengdu Institute
of Biology, Chinese Academy of Sciences (CIB); Kunming Institute of
Zoology, Chinese Academy of Sciences (KIZ), Kunming, Yunnan, China;
University of Kansas Biodiversity Institute (KU), Lawrence, KS, USA;
Field Museum of Natural Histor y (FMNH), Chicago, IL , USA; Museum
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of Comparative Zoology (MCZ), Cambridge, MA, USA; Natural History
Museum (NHM), London, UK; and Smithsonian National Museum of
Natural History (USNM).
2.2 | Morphological data and statistical analyses
Morphometric measurements and pholidosis data were taken from a
subset of the examined specimens (Tables S2 and S3). Morphometric
data were taken with a digital slide caliper by KW to the nearest
0.1 mm. Focal characters and character definitions follow Wang, Wu,
Jiang, et al. (2019), incl ud in g sn out–ve nt leng th (SV L); tai l le ng t h (TA L);
head width (HW ); he a d le n g th (H L) ; he a d de p t h (HD); s n o u t– e ye leng t h
(SEL); length of tallest nuchal crest (TNC); fore-limb length (FLL); hind
limb length (HLL); Toe IV length (T4L); trunk length (TRL); supralabial
count (SL); infralabial count (IL); number of scales between nasal and
first supralabials (NSL); number of scale rows between supralabials
and orbit circle (SOR); enlarged, modified, post-occipital scale count
(POS); enlarged, modified, post-tympanic scale count (PTY); enlarged,
modified, post-rictal scale count (PRS); Finger IV subdigital lamellae
count (F4S); Toe IV subdigital lamellae count (T4S); middorsal scale
count (MD); and keel status of ventral body scales (KVS). Values for
paired pholidosis characters (SL, IL, NSL, SOR, F4S, T4S, POS, PTY,
and PRS) were recorded from both side s of th e bo dy, with counts pro -
vided in lef t/right order, while for paired morphometric data (FLL and
HLL), a single value was taken at the left side.
Because coloration and ornamentation are both diagnostic (Wang,
Jiang, Ren, et al., 2019; Wang et al., 2016, 2017; Wang, Ren, Jiang,
et al., 2019; Wang, Wu, Jiang, et al., 2019) and play import ant roles
in the sexual selection processes for agamid lizards (Stuart-Fox &
Ord, 2004), data of coloration and ornamentation were also collected
from both live and preserved specimens, including (a) presence or ab-
sence of gular spot; (b) coloration of gular spot, if present; (c) presence
or absence of dorsolateral stripes on the body; (d) shape of dorsolat-
eral stripe of body (strongly jagged, feebly jagged, or smooth-edged),
if present; (e) coloration of dorsolateral stripes of body, if present;
(f) shape of vermiculated stripes on ventral head (reticulated versus
scattered); (g) coloration of inner lips; (h) coloration of oral cavity and
tongue; and (i) coloration of ventrolateral body. Color descriptions
were standardized using terminology and codes in Köhler (2012).
To account for the allometry between adult s and juveniles, which
is known in the genus Diploderma (Wang et al., 2015, 2016, 2017),
data of juvenile and subadult specimens were excluded from com-
parisons and morphometric analyses. Principal component analysis
(PCA) and discriminant analysis of principal components (DAPC)
were performed to examine whether the questionable populations
of D. cf. flaviceps from Yunn an an d Si chu an Province s occupy dist inc t
morphological spaces with respect to true D. flaviceps a n d ot her mor-
phologically similar congeners. These combined analyses allowed us
to determine whether the morphological clustering coincided with
the clades recovered from our molecular data set.
Because species of the genus Diploderma are known to be sexu-
ally dimorphic in morphometric measurements (Manthey et al., 2012;
Wang et al., 2015, 2018; Wang, Jiang, Ren, et al. , 2019), morpho metric
data of each sex are treated separately for analyses and comparisons.
Due to a limited sample size for D. drukdaypo, only females were in-
cluded for the morphometric analyses (males n < 3). Raw morpho-
metric measurements were used for statistical analyses, while relative
ratios of raw measurements (i.e., TAL/SVL, FLL/SVL, HLL/SVL, TRL/
SVL , HW/SVL, HD/SVL , HL/SVL, HW/HL, HD/H L, HD/HW, and SEL/
HL) were used for comparisons among species, in order to remove co-
variance among raw morphological measurements.
On the other hand, as scale counts are less prone to sexual dimor-
phism and allometry (Manthey et al., 2012; Wang, Jiang, Ren, et al., 2019;
Wang et al., 2016; Wang, Wu, Jiang, et al., 2019), pholidosis data of both
sexes and juveniles/subadults were combined in analyses and com-
parisons. A total of nine pholidosis counts (SL , IL, SOR, F4S, T4S, MD,
POS, PTY, and PRS) were used for the analyses. For each of the paired
counts (SL, IL, SOR , F4S, T4S, POS, PTY, and PRS), the mean values of
counts from both sides of the body of each individual were calculated
and rounded to the nearest whole numbers for the statistical analyses.
All morphological data were square root transformed prior to the
statistical analyses and scaled to their standard deviation in order to
normalize their distribution. PCA was per formed using the prcomp
command in R v. 3.2.1. Major PCs that explain at least 95% of the
total variance and the major discriminant functions were retained
and used for DAPC analyses, also using R v. 3. 2.1. The first two major
principal components from PC A and the results of DAPC analyses
were plotted using ggplot2 in R with 95% confidence ellipses.
Furthermore, one-way analysis of variance (ANOVA) was used to
search for the signific ant variation in morphometric data among the
putative new species and their morphologically similar congeners;
and if ANOVA yields significant variation, then Tukey's HSD test was
used to determine whether the putative new species are statistical
different from each of the given congeners. Both statistical test s
were done using R.
2.3 | Molecular data and phylogenetic analyses
Sequence data of 69 individuals were generated in this present study
and deposited in GenBank (Accession numbers MT577892–578014
and MT659027–659119; Appendix 1). In addition, a total of 48 in-
dividuals of 18 species of Diploderma and seven species of three
outgroup agamid genera (Acanthosaura, Calotes, and Pseudocalotes)
were obtained from GenBank (Appendix 1). The outgroup selec-
tion followed the most recent phylogenetic studies of Draconinae
(Wang, Che, et al., 2019).
Genomic DNA was extracted from liver samples stored in 95%
ethanol using Fujita's guanidine thiocyanate protocol (Esselstyn,
Garcia, Saulog, & Heaney, 20 08). Fragments of a single mitochon-
drial (mt) DNA locus, NADH dehydrogenase subunit 2 gene (ND2),
and the nearby tRNAs (Trp, Ala, Asn), and three nuclear DNA loci,
namely brain-derived neurotrophic factor (BDNF), oocyte matura-
tion factor (CMOS) , and RNA fi ng erp rint pr ot ein 35 (R35) were ampl i-
fied using published primers and PCR protocols (Crottini et al., 2009;
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WANG et Al .
Wang, Che, et al., 2019; Whiting, Bauer, & Sites, 2003; Table 1), and
the PCR products are about 1,200 bp, 200 bp, 600 bp, 500 bp, and
800 bp, respectively. After sequencing and trimming, the alignments
of new sequences are 1,032 bp, 184 bp, 567 bp, 391 bp, and 641
bp for ND2, tRNAs, BDNF, CMOS, and R35, respectively. When in-
cluded and aligned with the GenBank data, the final alignments are
1,032 bp, 392 bp, 567 bp, 539 bp, and 641 bp for the five genes,
respectively. Amplified products were visualized on 1.5% agarose
gels, and clean single banded products were purified with 1 μL of
a 20% dilution of ExoSAP-IT (US78201, Amersham Biosciences).
Cycle sequencing reactions were carried out using ABI Prism BigDye
Terminator chemistr y (Ver. 3.1; Applied Biosystems), and purified
with Sephadex (NC9406038, Amersham Biosciences) in Centri-Sep
96 spin plates (CS-961, Princeton Separations). Purified products
were sent to Eurofins Genomics for analysis.
Raw sequences were assembled and aligned using Geneious
v.10.0 with minor subsequent manual adjustments. The final align-
men ts and resul ting topologi es are deposite d in Dat a Dr yad (https://
doi.org/10.5061/dryad.7sqv9 s4qc.). Pairwise genetic distances of
the ND2 coding region among species of the genus Diploderma were
calculated using PAUP v. 4.0 b10 (Swofford, 20 02).
The phylogenetic trees were generated using both maxi-
mum-likelihood and Bayesian analyses. To assess phylogenetic con-
gruence between the mitochondrial and nuclear data, we inferred
the phylogeny for each gene independently using likelihood and
Bayesian analyses. As no strongly supported incongruences be-
tween mitochondrial and nuclear data were observed visually, we
concatenated the two data sets for the final analyses.
Partitioned Bayesian analyses were conducted using the pro-
gram MRBAYES v. 3.2.7a on CIPRES (Ronquist et al., 2012). All pro-
tein-coding genes were partitioned by codon positions, except for
the mitochondrial tRNAs, which were combined and treated as a
single partition (Table 1). The best model of nucleotide substitution
was selected for each par tition by the Akaike information criterion
(AIC), implemented in JMODELTEST2 v. 2.1.10 (Darriba, Taboada,
Doallo, & Posada, 2012; Guindon & Gascuel, 2003). A rate mul-
tiplier model was used to allow substitution rates to vary among
subsets. Two independent Markov chain Monte Carlo analyses
were run, each with four Metropolis-coupled chains. Because the
analyses did not converge under default settings, we adjusted the
melting temperature (from 0.2 to 0.1). All Bayesian analyses were
run for 2,000,000,000 generations, with parameters and topol-
ogies sampled every 2,000 generations. Stationarity and conver-
gence were assessed with TRACER v. 1.6.0 (Rambaut, Suchard, &
Drummond, 2013). We conservatively discarded the first 20% of
samples as burn-in.
Partitioned maximum-likelihood analyses were performed using
RAxML-VI-HPC v. 8.2.10 (Stamatakis, 2014) on the final data set
using the same partition strategy as for Bayesian analyses. The most
complex model (GTR + Γ) was applied for all subsets, with 1,000
replicate ML inferences run. Each inference was initiated with a
random starting tree, and nodal suppor t was assessed with 1,000
bootstrap pseudoreplicates. Nodes having ML bootstrap values of
70 and above and BI posterior probabilities of 0.95 and above were
considered well-supported.
3 | RESULTS
3.1 | Molecular results
The topologies of the consensus trees obtained from the ML and
BI analyses were very similar, except for three nodes (within the
outgroup genus Pseudocalotes, the node inside of node E, and node
Q), which either have low suppor t in the ML analyses and are not
resolved in the BI analyses (for the first and last cases) or differ-
ent topologies (regarding the relationships between D. swinhonis and
D. polygonatum; Figure 2). Tip nodes that unif y recognized species
are all well-supported (bootstrap support > 90, Bayesian posterior
probability > 0.95, herein all given in such order).
TABLE 1 Partition strategies, primers, and corresponding annealing temperatures for PCR of all genetic markers used in this study
Gene Codon Substitution model Primer name Primer info References
BDNF 1GTR BDN F. F CCCCA ATGAAAGAAGTGASCCTC Crottini et al. (2009)
2HKY + ΓB D NF.R TG GGTAGT TCGGCACTGAGA ATTCC
3JC69
CMOS 1JC69 G73.1 GGCTRTAAARCARGTGAAGAAA Whiting et al. (2003)
2HKY + ΓG 74.1 GARC WTCCAAAG TCTCC AATC
3HKY + Γ
R35 1 GTR + ΓR35.F GACTGTGGAYGAYCTGATCAGTGTGG Whiting et al. (2003)
2HKY R35.R GCCAAAATGAGSGAGAARCGCTTCTG
3GTR + Γ
ND2 1 GTR + ΓJap_70F CCACCA AAC AACTACACCTA Wang, Che, et al. (2019)
2GTR + Γ
3GTR + ΓJap_1559R GGAT TA ATG CC C TC TGG AT T
tRNAs 1 GTR + Γ
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WANG et Al.
The genus Diploderma is recovered as monophyletic (100/1.00).
Both D. drukdaypo and D. iadinum are recovered as members of the
genus Diploderma, as morphological data suggested. Members of the
D. flaviceps complex are paraphyletic, as members of the complex are
scattered across the genus (Figure 2). Within the genus Diploderma,
two ma j o r clade s, Cla de A (10 0/1 .0 0 ) and Cla de B (7 8 /0 .96) , ar e reco v -
ered with strong support. For Clade A, two major subclades, Clade C
and Clade F, are recovered. In Clade C, true D. flaviceps i s mo st clo se ly
related to D. splendidum (100/1.00), and the entire group is sister to
Clad e E, wh ic h co nta ins all specie s from Ea st As ian isl an ds (10 0/1 .00) .
Within Clade F, D. zhaoermii and D. micangshanense, D. varcoae and
D. dymondi, and D. swild and the previously identified D. cf. flavi-
ceps from Miansha County, southern Sichuan Province (population
8 in Figure 1), each form a monophyletic group with strong support
(Clades G [100/1.00], I [99/1.00], and J [96/1.00], respectively).
All of the remaining populations of D. cf. flaviceps are recovered
within the second major clade, Clade B, and most populations are
paraphyletic with respect to each other, except the two populations
from Yajiang County and Shangri-La/Deqin County, which form
Clade J (Figure 2). Diploderma iadinum is recovered as the most basal
lineage in Clade B, which is followed by Clade K (100/1.00) that uni-
fies D. yunnanense and D. chapaense and Clade L (100/1.0 0) that uni-
fies D. laeviventre and D. slowinskii. The remaining recognized species
and all remaining populations of D. cf. flaviceps (populations 6, 7, 12,
14, and 15) are recovered within the next clade, Clade N (100/1.00).
Within Clade N, previously identified D. cf. flaviceps from Baiyu
County of Sichuan Province (population 12) is the most basal, which
is followed by D. batangense (Node O, 99/1.00). Although relation-
ships of D. yulongense and the populati on of D. cf. flaviceps from Muli
County, southern Sichuan Province (population 7) are not resolved
(Node Q, 48/–), the remaining three populations of D. cf. flaviceps
are all nested within the next more crown clade, Clade R (95/1.00),
where the population from Shangri-La County, nor thwest Yunnan
Province (population 15) is the most basal one, and populations from
Yalong County, Sichuan Province (population 6), and Deqin Count y,
Northwest Yunnan Province (population 14), form a monophyletic
group with strong support (Clade T, 92/1.00), sister to the Clade U
that unifies D. vela and D. drukdaypo (98/1.00; Figure 2).
The uncorrected pairwise distances are more than 12% be-
tween true D. flaviceps and members of the D. flaviceps complex,
and more than 5.7% among the six examined populations of D. cf.
flaviceps from Sichuan and Yunnan Provinces (Table S 4). The lowest
interspecific genetic distance is between D. drukdaypo and D. vela,
which is 2.4%–2.7%, while the highest is bet ween D. dymondi and
D. laeviventre, which is 20.5%–22.1%. In addition, two species show
high intraspecific genetic variation, namely D. splendidum (6.1%) and
D. chapaense (5.7%) (Table S4).
3.2 | Morphological results
For pholidosis data, the first eight principal components (PC) ex-
plain 96.5% of the total variance, in which the first four PCs have
eigenvalues greater than one. The four major PCs that have eigen-
values greater than one load most heavily on PRS, MD, PTY, and
SOR, respectively (Table S5). When plotting the first two PCs, most
populations of the D. cf. flaviceps are cl ustered togeth er, except pop-
ulation 8 from the middle Yalong River; and true D. flaviceps only
overlaps partially with the main cluster of D. cf. flaviceps (Figure 3).
Using the first eight PCs and three discriminant functions, although
DAPC analyses still cannot separate most of the examined D. cf. fla-
viceps populations from recognized congeners, they further separate
the population 8 of D. cf. flaviceps from the middle Yalong River and
true D. flaviceps from the main cluster of D. cf. flaviceps (Figure 3).
For morphometric data of males, the first three PCs explain
95.4% of the total variance, in which only the first PC has an ei-
genvalue greater than one ( Table S6). For the first PC , it loads most
heavily on SVL and HL , and the remaining two major PCs load most
heavily on TAL and FLL. When plotting the first two PCs, similar to
the PCA results of pholidosis data, most of the examined popula-
tions of D. cf. flaviceps are still clustered together, except population
12 from the far upper Jinsha River and population 8 from the mid-
dle Yalong River; and true D. flaviceps only overlap par tially with the
main cluster of D. c f. flaviceps (Figure 4). Using the first three PCs
and two discriminant functions, DAPC analyses further separates
the cluster, particularly between the true D. flaviceps and the six ex-
amined D. cf. flaviceps populations, and bet ween the main cluster
and population 8 along the middle Yalong River and population 12
from the far upper Jinsha River (Figure 4).
For morphometric data of females, the first four PCs explain
96.5% of the total variance, in which the first two PCs have eigenval-
ues greater than one (Table S7). For the first two PCs, they load most
heavily on HD, HL, TAL, and TRL. Unlike the pattern for the males,
PCA plots of the first two PCs show two major clusters for females:
One contains four of the six examined populations of D. cf. flaviceps,
true D. flaviceps, and three of the included recognized species of
the complex (i.e., D. batangense, D. swild, and D. yulongense), while
the other one cluster consist s of two population of D. cf. flaviceps
(populations 6 and 12), as well as D. drukdaypo (Figure 4). Using the
first four PCs and two of the discriminant functions, DAPC analyses
further separate the clusters, including the true D. flaciceps from the
main cluster (Figure 4).
Despite the overlapping clustering among many of the examined
D. cf. flaviceps populations in morphospace, when comparing color-
ation/ornamentation (i.e., shape and color of dorsolateral stripes,
gular coloration, ventrolateral coloration, and oral coloration) and
development statuses of pholidosis characters (i.e., degree of modi-
fication of conical scales on the head and crest shape), D. cf. flaviceps
populations that overlap in morphospace of morphometric and pho-
lidosis characters can be readily distinguished from each other and
from all recognized congeners (see comparison sections below for
details; Tables 2–4; Figures 2, 11, 12).
In conclusion, the six examined populations of D. cf. flaviceps
from Yunnan and Sichuan Provinces are similar to members of
the D. flaviceps complex in terms of morphometric and pholidosis
characters, but each of them can still be diagnosed readily from all
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WANG et Al.
recognized species particularly using coloration and ornamentation
cha ra cter s. In addition , the genealog y from th e mu lti-l oc us dat a se t
shows that each of the six populations also forms a monophyletic
group, which is paraphyletic to true D. flaviceps. Fur thermore, each
of the six populations also exhibits considerable genetic distance
from all other recognized congeners and from each other (>5.7%).
Therefore, adopting the evolutionar y species concept, we here
consider each of the six examined D. cf. flaviceps populations as
distinc t new species.
3.3 | Taxonomic accounts
3.3.1 | Diploderma angustelinea sp. nov. Wang, Ren,
Wu, Che, Siler
(Population 7 in Figure 1; Figures 2, 5, 11, 12;
Tables 2–4; Table S8 (1); ZooBank ID urn:lsid:zoobank.
org:act:E709207E-3FC6-45EF-9D54-8CADAE8949A8).
Synonym
Japalura flaviceps Zhao et al. (1999: 111–115), in part; Zhao (2003:
84), in part.
Holotype
KIZ 029703, adult male from Maidilong Village, Muli Tibetan
Autonomous County, Liangshan Prefecture, Sichuan Province, China
(28.594° N, 101.226° E, elevation 2017 m, WGS 84). Collected by
locals on June 10, 2016.
Paratopotypes
KIZ 0 44 484, 044796, 044797, adult males; collected by Kai Wang,
Zhuoyu Lu, and Xiankun Huang on June 25, 2019. KIZ 029704–
029708, 029710, adult females, same collecting information as for
the holot ype.
Diagnosis
The new species can be diagnosed from congeners by a combination
of the following morphological characteristics: (1) body size moderate,
SVL 56.8–60.2 mm in males, 59.0–67.8 mm in females; (2) tail long,
TAL 230.2%–249.1% SVL in males, 194.30%–222.3% in females; (3)
hind limbs moderate, HLL 71.1%–80.2% SVL in males, 74.4%–79.2% in
females; (4) head width moderate, HW 66.6%–73.9% HL, HD 72.7%–
79.3% HW; (5) MD 41–49; (6) F4S 14–19; (7) T4S 21–26; (8) post-rictal
sub-pyramidal scales weakly developed and few, 0–3; (9) tympanum
co nce ale d; (10 ) nu c hal cre s t sca l es weak ly dev e lop ed wi tho u t sk i n fo l ds
in both sexes, slightly raised and serrated in males; (11) distinct trans-
verse gular fold present, deep; (12) ventral scales of head and body
distinc tively keeled; (13) gular spots always present in males, mostly in
females, Spectrum Yellow (Color 79) to Dark Spectrum Yellow (Color
78) in lif e, abs en t af te r pr es er vation; (14) no dark vermiculate stripe s on
ventral head; (15) dorsolateral stripes narrow, feebly jagged, Spectrum
Yellow (Color 79), present in both sexes; (16) dark radial stripes around
eye s in disti nc t or absent ; (17) dis tin ct, cle ar sep arati on bet wee n th e Jet
FIGURE 3 PCA (1) and DAPC plots (2) with 95% confidence ellipses based on nine pholidosis characters (SL, IL, SOR, F4S, T4S, MD, POS,
PTY, and PRS) for the six examined populations of D. cf. flaviceps and their morphologically similar congeners. The embedded figure in the
DAPC plot indicates the DA eigenvalue for each of the major discriminant functions, where the shaded ones were obt ained for the DAPC
analyses. See methods for abbreviations and their morphological definitions
FIGURE 2 Phylogenetic relationships of the genus Diploderma inferred from a concatenated multi-locus data set (ND2, RNAs, BDNF,
CMOS, and R35), with gular patterns of males and females in life of the D. flaviceps complex shown on the side. All tip nodes that unify each
individual species arewell-supported (BS > 90, BP > 0.98). “–” indicates non-resolved or differential relationship in Bayesian analyses. Shaded
species are members of the D. flaviceps complex, while new species described here are shaded with the same color as they appear in the
map in Figure 1. The circled number of each species corresponds to the number shown in Figure 1. Photographs by Kai WANG and Christian
WEINERT
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WANG et Al .
Black (Color 300) or Burnt Umber (Color 48) coloration of dorsolateral
body and white coloration of ventrolateral body; and (18) oral cavity
and tongue Light Flesh Color (Color 250).
Comparisons
Diploderma angustelinea sp. nov. was confused historically with
D. flaviceps, with both species having faint, indistinct, or no dark ra-
dial stripes around eyes. However, the new species can be distin-
guished from the latter by having feebly developed nuchal crests
without skin folds in both sexes (versus well-developed on skin folds
in both sexes), a relatively longer tail in males ( TAL 230. 2%–249.1%
SVL versus 181.8%–210.8% SVL ), fewer and weakl y develope d po st-
rictal modified scales (0–3, average 1, sub-pyramidal-shaped and
weakly developed versus 3–9, average 6, conical-shaped and well-
developed), feebly jagged, narrow dorsolateral stripes in both sexes
(versus strongly jagged and wide), a sharp transition in body colora-
tion from dorsolateral Jet Black (Color 300) or Burnt Umber (Color
48) coloration to ventral white (versus gradual transition from Pale
Pinkish Buf f [Color 3] to white), and by the presence of distinct gular
spots in both sexes (versus absence), and absence of reticulated, ver-
miculate black patterns on the gular region (versus presence).
The new species is morphologically most similar to D. laeviven-
tre, in which both species possess yellowish gular spots, distinct
transverse gular folds, and none or few weakly developed post-rictal
modified scales. However, D. angustelinea sp. nov. can be differen-
tiated from the latter species by having distinctively keeled ventral
scales (versus feebly keeled or smooth), fewer middorsal scales
(41–49 versus 57–59), slightly jagged, narrower dorsolateral stripes
(versus completely smooth-edged and wide), and by the absence of
heavy black speckles on dorsal, lateral, and ventral surfaces of head
(versus presence).
Diploderma angustelinea sp. nov. differs from all remaining
recognized congeners by having narrow, dorsolateral stripes in
both sexes (versus thick and wide). Specifically, D. angustelinea
FIGURE 4 PCA (1) and DAPC plots (2) with 95% confidence ellipses based on nine morphometric characters (SVL, TAL, HL , HW, HD,
SEL, FLL , HLL, and TRL) for both males (a) and females (b) of the examined populations of D. cf. flaviceps and their morphologically similar
congeners. The embedded figure in the DAPC plot indicates the DA eigenvalue for each of the major discriminant functions, where the
shaded ones were obt ained for the DAPC analyses. See methods for abbreviations and their morphological definitions. For population 6 and
population 14, as only female and male specimens were collected from each population, respectively, the analyses for each sex only have
representation from five D. cf. flaviceps populations in total
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TABLE 2 Comparisons of morphometric data among the six new species described here (Diploderma angustelinea sp. nov., D. aorun sp. nov., D. flavilabre sp. nov., D. panlong sp. nov., D. qilin
sp. nov., and D. panchi sp. nov.) and their morphologically similar congeners (D. batangense, D. drukdaypo, D. flaviceps, D. swild, D. vela, and D. yulongense). See methods for abbreviations and
their morphological definitions. Average values are given in parentheses
Species Sex
Sample
size SVL TAL/SVL (%) FLL/SVL (%) HLL/SVL (%) HL/SVL (%) HD/HW (%) HW/HL (%)
D. angustelinea sp.
nov.
M 4 56.8–60.2 (58.5) 230.2–249.1 (235.3) 45.7–46.5 (46.4) 71.1–80.2 (77.0) 27.8–30.1 (29.4) 73.3–78.7 (76.4) 66.6–70.8 (68.7)
F659.0–67.8 (63.1) 194.3–222.3 (207.3) 40.8–4 6.2 (43.7) 74.4–79.2 (75.9) 2 7.3–2 9.3 ( 28. 2) 72.7–79.3 (75.9) 67.0–73.9 (68 .9)
D. aorun sp. n ov. M756.3–61.2 (58.7) 212.0–221.1 (216.4) 43.7–50.4 (47.7) 73.7–81.7 (78.6) 2 9.2–3 1.4 (2 9.9) 71.4–8 0.2 (75.4) 67.8–72.7 (70.8)
F 4 60. 8–66.5 (63.8) 191 .4 –207.7 (19 9.2) 45.0–52.8 (47.9) 71.5–80.5 (75.3) 27.3–30. 2 (28.9) 66.7–78.0 (74.0) 66.8–70.4 (68.7)
D. flavilabre sp.
nov.
M350.8–55.2 (53. 5) 149.7–179.3 (176.0) 43.1–48.3 (45.8) 66.9–74.8 (71 .5) 31.2–32.2 (31.6) 70.1–75.5 (72.4) 70.7–73.1 (71.6)
F664.2–72.9 (67.5) 140.2–152.2 (146.4) 40.1–43.5 (41.2) 62.0–65.6 (63.0) 27.1–29. 3 (2 8 .6) 68 .6–75.1 (71.3) 70.1–74.9 (71. 8)
D. panchi sp. nov. F 4 59.6–67.7 (65.2) 141.8–151.5 (148.1) 39.3–41.7 (40.0) 60.2–65.8 (62.6) 27.0–28.8 (28.0) 63.3–74.3 (69.0) 69.5–73.6 (71.6)
D. panlong sp. n ov. M 4 64.2–71.7 (64.8) 248.2–268.0 (257.1) 44.9–46.9 (46.3) 77.7–83.2 (80.5) 29.1–31.6 (30.7) 77.8–79.5 (78.5) 63.4–67.3 (65.0)
D. qilin sp. n ov. M755.9–66. 5 (55.9) 201.0 –218.2 (20 8.6) 44.3–48.5 (46.4) 73.3–78.4 (75.8) 28.8–31.2 (31. 2) 73 .9–82.7 (80.4) 65.0–72.7 (68.1)
F655.3–72.0 (61.0) 174.3–199.7 (187.5 ) 44.7–51.8 (46.8) 72.5–80.4 ( 74.5) 26.6–28.9 (28.0) 72.3–80.4 (76.2) 69.6–73.2 (71.1)
D. batangense M655.5–62.8 (59.3) 187.3–206. 5 (194.6) 46.0–49.8 (47.9) 74.7–79.8 ( 76.4) 28.8–32.2 (30.9) 71.7–75.3 (73.6) 66 . 3 –67. 3 (67.0)
F 5 54.7–63.1 (60.0) 171.8–203.1 (186.9) 46.2–51.2 (48.4) 73.4–79.2 (76.9) 28.5–31.7 (30.2) 68.4–75.9 (72.5) 65.3–71.5 (68.1)
D. drukdaypo M249.9–51.5 (50.7) 153.1–154.4 (153.7) 37.9–45.4 (41.6) 58.2–63.8 (61.0) 27.2–33.2 (30. 2) 71.1–73.2 (72.1) 65.9–66.7 (66.3)
F650.2–58.9 (54.3) 132.8–143.9 (138.0) 37.5–42.6 (39.4) 59.7–62.8 (60.8) 27.8–30.2 (28.9) 66.3–73.8 (68.7) 66.4–72.1 (70.5)
D. flaviceps M 5 68.5–82.1 (74.9) 181.8–210.8 (191.6) 42.3–45.6 (44.2) 69.7–73.5 (71.9) 30.7–32.7 (31.9) 76.7–85.1 (81.2) 63.1–65.4 (64.4)
F359. 6– 69.3 ( 6 4.2) 191.5–205.7 (197.0) 44.7–46.0 (45.4) 69.3–77.8 (73.7) 31.6–31.8 (31.7) 75.1–81.0 (78.3) 64.5–67.9 (66.5)
D. swild M359.8–71.7 (66.6) 2 24.4–2 39. 0 (233 .3) 46.3–48.7 (47.2) 72.0–78.1 (75.1) 31.2–32.5 (32.0) 73.4–83.4 (78.6) 65.8–71.5 (67.9)
F 4 59.7–76.8 (68.8) 200.4–221.0 (208.8) 41.9–49.2 (44.9) 69.4–75.9 (70.7) 30.8–34.6 (32.1) 77.0–80.6 (78.5) 64.9–70.0 (67.2)
D. vela M10 51.6–64.5 (58.2) 174 .6–238 .1 (19 9.3) 43.5–53.1 (47.9) 67. 5–85.6 (75.6) 30.5–32.8 (31.4) 66.7–76.7 (73.3) 64.5–69.3 (67.8)
F10 54.1–64.2 (59.3) 159.8–202.2 (180.0) 43. 3–51.8 (47.0) 65.7–81.6 (72.9) 28.2–30.4 (29.7) 67.6–73.8 (70.7) 65.2–69.8 (67.8)
D. yulongense M 9 56.2–70.7 (62.9) 193.7–244.3 (223.0) 42.9–47.9 (45.1) 69.1–78.0 (74.4) 29.3–32.3 (30.7) 77.9–88.3 (82.4) 62.1–72.4 (67.2)
F656.2–69.2 (62.5) 182.2–211.3 (194.4) 42.0–45. 3 (44.0) 68.8–74.9 (71.2) 27.8–30.2 (29.1) 73.4–83.8 (78.9) 63.7–68.5 (66.4)
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differs from D. brevicaudum and D. drukdaypo by having a longer
tail (230.2%–249.1% SVL in males, 194.3%–222.3% SVL in females
versus 140.0% in male, 125.0%–145.0% in females for D. brevi-
caudum; 153.0%–154.4% in males, 132.8%–144.0% in females for
D. drukdaypo) and much longer hind limbs (HLL 71.1%–80.2% ver-
sus 60.0%–64.0% in D. brecicaudum, 58.2%–63.8% in D. drukdaypo);
from D. batangense, D. iadinum, D. vela, D. yulongense, and D. zhao-
ermii by having weakly developed nuchal crests without skin folds
(ver sus wel l-developed on rais ed on skin fo lds in male s) an d distinc t
coloration of gular spots (Dark Spectrum Yellow [Color 78] ver-
sus Pale Cyan [Color 157] in D. batangense; Caribbean Blue [Color
168] and Medium Greenish Yellow [Color 88] males and females
in D. iadinum, respectively; Chartreuse [Color 89] in D. yulongense
and D. zhaoermii; and no gular spots in D. vela); from D. chapaenses,
D. micangshanense, D. varcoae, D. yunnanenses, and all island species
(D. brevipes, D. luei, D. makii, D. polygonatum, and D. swinhonis) by the
presence of a distinct, deep, transverse gular fold (versus absence);
from D. fasciatum by the feebly developed nuchal crests (versus
well-developed and differentiated from dorsal crests), and by the
presence of dorsolateral stripes (versus absence) and the absence
of hourglass-shaped pattern on the mid-dorsum (versus presence);
from D. dymondi, D. slowinskii, D. swild, and D. varcoae by having a
concealed tympanum (versus exposed); from D. splendidum by hav-
ing a smaller maximum body size (SVL ≤ 67.8 mm versus ≤92.0 mm),
homogeneous ventral head scales (versus heterogeneous), and dis-
tinct, deep, transverse gular fold (versus shallow); from D. hamptoni
by having parallel dorsolateral stripes (versus diagonally away from
vertebral line posteriorly), a distinct transverse gular fold (versus
shallow), and by the absence of distinct dark stripes on gular region
(versus presence).
Description of holotype
Medium-sized agamid, SVL 57.6 mm, body slender, not dorsally
compressed; tail slender, long, TAL 230.2% SVL; limbs moderate
in length, FLL 45.8% SVL , HLL 79.1% SVL; head robust, HW 70.6%
HL, HD 75.84% HW, 53.5% HL; snout pointy, SEL 36.9% HL. Rostral
rectangular, four times wider than high, one or two small scales in
direct contact between rostral and nasal; nasal oval, naris positioned
mediodorsally; supralabials 9/9, anteriormost supralabial single scale
away from nasal on both sides, posteriormost longest, approxi-
mately three times longer than others, each bearing single, weak,
lateral keel; infralabials 9/9, posteriormost shortest, each bearing
sin gl e, late ral ke el toward inferior sid e; conic al or subpy ramida l po st-
rictal scales, 2/2, relatively weak; suborbital scale rows 4/4, sube-
qual in sizes; enlarged, elongated, keeled scales between posterior
orbit and anterosuperior tympanum, 7/7; tympana concealed under
fine scales; post-tympanic conical scales well-developed, protruding,
2/5.
Dorsal head scales keeled, heterogeneous in size and shape; sin-
gle Y-shaped ridge formed by series of hexagon-shaped, enlarged
scales on dorsal snout, with end of Y-shaped ridge two small scales
posterior to rostral, tips of Y-shaped ridge consisting of single pair
of distinctively enlarged, protruding scales, lining up at line between
anterior orbits; supraciliaries 7/7, overlapping more than one third of
total length with neighboring scales; enlarged, somewhat elongated,
oval-shaped, protruding scale dorsoposterior to orbit, 1/1; interpari-
etal enlarged, irregular-shaped, with distinct parietal eye; two lateral
series of four enlarged, slightly protruding scales symmetrical along
vertebral axis posterior to interparietal on each side of occipital
head; enlarged, conic al scales on post-occipital head, 4/6, dorsally
fattened, pointing posteriorly.
TABLE 3 Comparisons of pholidosis data among the six new species described here (Diploderma angustelinea sp. nov., D. aorun sp. nov.,
D. flavilabre sp. nov., D. panlong sp. nov., D. qilin sp. nov., and D. panchi sp. nov.) and their morphologically similar congeners (D. batangense,
D. drukdaypo, D. flaviceps, D. swild, D. vela, and D. yulongense). See methods for abbreviations and their morphological definitions. Average
values are given in parentheses. For keel status of ventral scale, S: strongly keeled; N: not keeled; W: weakly keeled
Species
Sample
Size F4S T4S MD PTS PTY PRS KVS
D. angustelinea sp. nov. 10 14–19 (17) 21–26 (23) 41–49 (45) 1–5 (4) 1–4 (2) 0–3 (1) S
D. aorun sp. n ov. 11 12–18 (15) 16–24 (21) 35–46 (40) 1–4 (3) 2–6 (4) 1–7 (4) S
D. flavilabre sp. n ov. 9 13–16 (15) 19–22 (21) 39–43 (41) 3–6 (4) 3–7 (5) 4–9 (6) S
D. panchi sp. nov. 414–17 (15) 20–24 (22) 42–46 (45) 2–5 (3) 2–6 (4) 3–7 (4) S
D. panlong sp. n ov. 718–24 (20) 22–27 (25) 39–46 (42) 2–6 (4) 6–12 (9) 9–13 (11) S
D. qilin sp. n ov. 13 15–19 (17) 21–25 (22) 38–45 (41) 2–5 (4) 3–8 (6) 3–8 (5) S
D. batangense 11 13–17 (15) 18–24 (21) 41–53 (44) 2–5 (3) 1–5 (3) 0–3 (2) S
D. drukdaypo 813–17 (15) 18–23 (20) 43–56 (48) 1–5 (3) 1–4 (2) 0–3 (2) N or W
D. dymondi 12 16–21 (19) 22–27 (24) 41–51 (45) 2–5 (3) 4–9 (6) 6–10 (9) S
D. flaviceps 11 15–22 (18) 22–27 (24) 45–56 (51) 2–6 (4) 4–8 (6) 3–9 (6) S
D. swild 9 18–22 (20) 23–27 (25) 35–44 (42) 1–3 (2) 3–7 (5) 5–10 (8) S
D. vela 28 14–18 (16) 20–25 (22) 40–50 (44) 2–5 (4) 1–5 (3) 0–6 (3) S
D. yulongense 14 15–19 (17) 21–26 (24) 38–4 4 (40) 1–5 (3) 1–3 (2) 0–4 (2) S
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TABLE 4 Comparisons of coloration and ornamentation data among the six new species described here (Diploderma angustelinea sp. nov., D. aorun sp. nov., D. flavilabre sp. nov., D. panlong
sp. nov., D. qilin sp. nov., and D. panchi sp. nov.) and their morphologically similar congeners (D. batangense, D. drukdaypo, D. flaviceps, D. swild, D. vela, and D. yulongense). Abbreviations see
methods. “+”: present; “−”: absent; SFNC: Skin Fold under Nuchal Crest; SFDC: Skin Fold under Dorsal Crest; CSSE: Crest Scale Strongly Erected;. For color codes see Köhler (2012)
Species Sex SFNC SFDC CSSE
Inner-Lip
Coloration Tongue Coloration
Pharyngea
and Palatum
Coloration
Gular
Spot
Gular Spot
Coloration
Dorsolateral
Stripe Shape
Ventral Body
Coloration
Ventrolateral
Body Coloration
D. angustelinea
sp. nov.
M − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Dark Spectrum
Yellow (78)
Feebly jagged White White or Light
Buff (Color 1)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Dark Spectrum
Yellow (78)
Feebly jagged White White or Light
Buff (Color 1)
D. aorun sp.
nov.
M+ + + Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Light Cyan
(159)
Strongly jagged White or Pale
Sulphur
Yel lo w
(Color 79)
White or Light
Buff (Color 1)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Light Cyan
(159)
Strongly jagged White White or Light
Buff (Color 1)
D. flavilabre
sp. nov.
M+ + + Spectrum Yellow Light Flesh Color
(250)
Light Flesh Color
(250) speckled
with Pale Neutral
Gray (296)
+Pale Emerald
Green (141)
to Light
Tur quo ise
Green (146)
Strongly jagged Pale
Greenish
Yel lo w
(Color 86)
Pale Buf f (1)
F − − +Spec trum Yellow Light Flesh Color
(250)
Light Flesh Color
(250) speckled
with Pale Neutral
Gray (296)
+Pale Emerald
Green (141)
to Light
Tur quo ise
Green (146)
Strongly jagged White White or Light
Buff (Color 1)
D. panchi sp.
nov.
F − − − Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
D. panlong sp.
nov.
M − − +Dark Spectrum
Yellow (78)
Dark Spectrum
Yellow (78)
Dark Spectrum
Yellow (78)
− − Smooth White White
D. qilin sp. n ov. M+ + + Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Light Sulphur
Yellow (Color
93)
Strongly jagged White Light Sulphur
Yellow (93)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Light Sulphur
Yellow (Color
93)
Strongly jagged White White
D. batangense M+ + + Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Pale Cyan
(157)
Strongly jagged Sulphur
Yel lo w
(Color 79)
Sulphur Yellow
(Color 79)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Pale Cyan
(157)
Strongly jagged White White or Light
Buff (Color 1)
(Continues)
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WANG et Al .
Species Sex SFNC SFDC CSSE
Inner-Lip
Coloration Tongue Coloration
Pharyngea
and Palatum
Coloration
Gular
Spot
Gular Spot
Coloration
Dorsolateral
Stripe Shape
Ventral Body
Coloration
Ventrolateral
Body Coloration
D. drukdaypo M − − − Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
F − − − Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
D. dymondi M − − +Smoky White
(261)
Light Flesh Color
(250)
Spect rum Violet
(186) to Jet Black
(300)
− − Smooth White White or Light
Buff (Color 1)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Spect rum Violet
(186) to Jet Black
(300)
− − Smooth White White or Light
Buff (Color 1)
D. flaviceps M+ + + Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
F+−+Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
D. swild M − − +Light Chrome
Orange (76)
Light Chrome
Orange (76)
Light Chrome
Orange (76)
− − Smooth White White or Light
Buff (Color 1)
F − − +Light Chrome
Orange (76)
Light Chrome
Orange (76)
Light Chrome
Orange (76)
− − Smooth White White or Light
Buff (Color 1)
D. vela M+ + + Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
− − Strongly jagged White White or Light
Buff (Color 1)
D. yulongense M+−+Smoky White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Chartreuse
(89) to
Opaline
Green (106)
Strongly jagged White Pale Green (99)
to Chartreuse
(89)
F − − +Smok y White
(261)
Light Flesh Color
(250)
Light Flesh Color
(250)
+Chartreuse
(89) to
Opaline
Green (106)
Strongly jagged White White or Light
Buff (Color 1)
TABLE 4 (Continued)
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WANG et Al.
Dorsal body scales heterogeneous in size and shape, all distinc-
tively keeled; axillary sc ales smaller than background dorsal scales,
non-granular in shape; enlarged scales two to four times larger than
surrounding background scales, some arranged in three parallel, dor-
solateral rows on each side of body: first row t wo scales inferior to
vertebral crest, second row through dorsolateral stripe, last row four
to five scales inferior to dorsolateral stripe; remaining enlarged dor-
sal scales scat tered randomly. Middorsal scale count 45; nuchal crest
scales serrated, slightly erect on weak skin fold in life (indistinct after
preser vation), short, CL 3.3% HL; dorsal crest scales relatively low,
enlarged, roughly equal in size to enlarged dorsal scales, serrated.
Dors al lim b sc al es dis tinct iv el y keeled , ho mo ge ne ou s in size on fore-
limbs, heterogeneous on hind limbs, with enlarged scales possessing
protruding keels on posterior lateral thigh. Finger I shortest, Finger
IV longest, F4S 16/16; Toe IV longest, T4S 24/26.
Ventral head scales distinctively keeled, mostly homogeneous
in size and shape, except three scales at center of gular pouch,
which are slightly enlarged; mental pentagonal in shape, sand-
wiched between first pair of chin shields; all chin shields except
first pair separated from infr alabials by one to three rows of small,
elongated scales; first two pairs of chin shields largest; gular pouch
well-developed in life, indistinct after preservation; transverse
gular fold present, deep, distinct. Scales on ventral body, limb, tail
mostly homogeneous in size, shape, distinctively keeled, regularly
arranged; keels of ventral tail scales carinate in lateral rows.
Coloration of holotype in life
The background color of the dorsal and lateral surfaces of the head
is uniform Medium Neutral Gray (Color 298), and the coloration is
much darker toward the occipital region of the head. Two transverse
bands occur between the eyes, which are more distinct toward the
terminal ends on both sides; and both bands are Robin Rufous (Color
29) on the terminal ends and gradually fade into Buff (Color 5) to-
ward the middle. Jet Black (Color 30 0) to Dark Neutral Gray (Color
299) radial stripes are present around the eyes, where most stripes
extend outside of the orbit circle except ones that are inferior to the
orbit. Radial stripes around the eyes are somewhat faint and indis-
tinct, except the posterior one that is toward the rictus on each side,
which is the broadest and most distinct stripe. A single Cream Color
(Color 12), distinct suborbit al stripe is present below the eye on each
side of the head, ex tending from the posterior nasal scale to the ric-
tus and beyond on each side. A somewhat faint, dark stripe, which
is formed by two rows of heavily speckled scales (Medium Neutral
Gray, Color 298), is present below the suborbital stripe from the
anteriormost supralabial to the second most posterior supralabial
scale. Supralabial scales are Light Yellow Ocher (Color 13), distinct
FIGURE 5 Male holotype (a1 and a2; KIZ 029703) and female paratopotype (b1 and b2 KIZ 029706) of Diploderma angustelinea sp. nov. in
life. Photographs by JR
(a1) (a2)
(b1) (b2)
236
|
WANG et Al .
from the infralabial scales. Background color on the post-tympanic
regions of the head is Smoky White (Color 261). Two short, longi-
tudinal, Dark Neutral Gray (Color 299) streaks (less than 10 scales
in leng th) are present on each side, which are parallel to each other
and positioned at the superior and inferior edges of the t ympanum,
respectively. For the remaining scales of the post-tympanic and
FIGURE 6 Male paratype (a1 and a2; KIZ 044764) and female paratype (b1 and b2 KIZ 032735) of Diploderma aorun sp. nov. in life.
Photographs by KW
(a1) (a2)
(b1) (b2)
FIGURE 7 Male holotype (a1–a3; KIZ 032693) and female paratopotype (b1–b3; KIZ 032695) of Diploderma flavilabre sp. nov. in life.
Photographs by KW
(a1) (a2) (a3)
(b1) (b2) (b3)
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237
WANG et Al.
post-rictal regions of the head, the keels and conical tips of all scales
are Sulphur Yellow (Color 80). The oral cavity and tongue are uni-
form Light Flesh (Color 250) colored in life.
Two narrow, Sulphur Yellow (Color 80) dorsolateral stripes are
present from the pos terior occipital region to the pelvis, one on each
side of the vertebral crest, parallel to each other. The dorsal region
between the two dorsolateral stripes is Orange-Rufous (Color 56),
with five rectangular, Burnt Umber (Color 48) patches scattered
evenly from the neck to the pelvis. Body sur faces inferior to the dor-
solateral stripes are Jet Black (Color 300) on the anterior part of the
body, fading into Burnt Umber (Color 48) and eventually to Drab-
Gray (Color 256) po sterior to the pelvis. Vent rolat er al sur fa ces of th e
bo d y are Sm oky Wh i te (C olo r 261) to whi te, wh ich is dis t inc t fro m the
blackish coloration below the dorsolateral stripes. Some of the en-
larged scales on the lateral body surfaces inferior to the dorsolateral
stripes are much lighter, which bear an either Flesh Ocher (Color 57)
or Sulphur Yellow (Color 80) medial keel. The dorsal surfaces of the
limbs are uniform Prat t's Payne's Gray (Color 293), with keels of in-
dividual scales in Olive Sulphur Yellow (Color 90). The dorsal and lat-
eral tail surfaces are Pale Neutral Gray (Color 296), with Flesh Ocher
(Color 57) scales scattered randomly near the tale base. Numerous
faint Light Flesh Color (Color 250) bands are scattered evenly on the
tail from one third of its length posteriorly, and the color eventually
fades into uniform Drab-Gray (Color 256) toward the end.
The background coloration of the ventral surface of the head is
white. A distinct, Spec trum Yellow (Color 79) gular spot is present on
the center of the gular pouch. Scales surrounding the gular spot bear
Sp ectr um Yel low (Co lor 79) kee ls . No di s ti n ct da rk ve rmi cul at e str ipe s
are present on the ventral surfaces of the head. The ventral surface
of the body is uniform white with no distinct color pigmentation, and
the ventral surfaces of the tail are uniform Smoky White (Color 261).
Coloration of holotype in preservation
Ornamentation patterns remain largely consistent after preserva-
tion; however, some observed coloration patterns fade, including the
Spectrum Yellow (Color 79) color of the gular spot and the Sulphur
Yellow (Color 80) color of the dorsolateral stripes turn to white, and
the Burnt Umber (Color 48) coloration of the dorsolateral regions
of the body become Dark Neutral Gray (Color 299) after long-term
preservation.
Variatio n
Morphometric and pholidosis variation of the type series
is summarized in Table S8. Although the sample size is lim-
ited, sexual dimorphism is evident in D. angustelinea sp. nov.,
with males differing from females by having a longer tail (TAL
230.2%–249.1% SVL versus 194.3%–222.3%), more developed
and serrated nuchal crests (versus less developed), and brighter
dorsolateral stripes (versus duller with less distinct edges), and
by the absence of distinct chevron patterns along dorsal mid-
line between dorsolateral stripes (versus presence). For some
females (KIZ 029704, 029708), the dorsolateral stripes are not
continuous , with three to four shor t breaks (one to four scales in
length) along the stripes, and a single female (KIZ 029704) lacks
the gular spot.
Etymology
The Latin specific name, “angustelinea”, comprises two parts: anguste
meaning “narrow” and linea meaning “stripe” or “line.” Together,
the specific name describes the diagnostic narrow, thin, dorsolat-
eral stripes present in the species. We recommend Narrow-striped
Mountain Dragon as its English common name and 细纹龙蜥 (Pinyin:
Xi Wen Long Xi) as its Chinese common name.
Natural Histor y and conservation status
The new species was found in arid grassy valleys along the mid-
dle Yalong River (Figure 13a). Individuals were observed to be ter-
restrial, basking and seeking shelters in rock crevices (sandstone)
during the day, but sleeping on bushes at night (i.e., Rumex hasta-
tus, the most common bush species in their habitats). Only a sin-
gle female was gravid (KIZ 029705), with all the remaining females
showing characters of recent oviposition (i.e., empty stomach and
extra skin folds on ventrolateral and lateral trunk), which suggests
that mid-June is toward the end of its breeding season. Currently,
the new species is known only from its type locality only in the
upper Yalong River Valley. Although no further information is
FIGURE 8 Female holot ype (KIZ 032715) of Diploderma panchi sp. nov. in life. Photographs by K W
(a1) (a2)
238
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WANG et Al .
available about its distribution range, rapid township expansion
and major hydropower developments were observed near the type
locality, which post major threats to the habitats of the species in
the hot–dry valleys. We recommend Data Deficient (DD) for its
IUCN classification, but call for future ecological and population
studies of this species.
3.3.2 | Diploderma aorun sp. nov. Wang, Jiang,
Zheng, Xie, Che, Siler
(Population 14 in Figure 1; Figures 2, 6, 11, 12; Tables 2–4;
Table S8 (2); ZooBank ID urn:lsid:zoobank.org:act:E81CAE00-
16CA-4D9A-8BD5-0B49158DFFF0).
Synonym
Japalura flaviceps Zhao and Yang (1997: 165–167), in part; Zhao
et al. (1999: 111–115), in part; Yang and Rao (2008: 200), in part.
Holotype
KIZ 044735, adult male, from Dari Village, Deqin County, Yunnan
Province, China (99.1733° E, 28.5804° N, elevation 2,198 m, WGS
84). Collec ted by Kai WANG, Xiankun HUANG, and Zhuoyu LU on
June 17, 2019.
Paratypes
KIZ 032734, 032736, 032737, adult males; KIZ 032735, adult fe-
male, all from areas near Benzilan township, Deqin County, Yunnan
Province, China (99.3827° E, 28.1774° N, elevation 2010 m, WGS
84); collected by Kai WANG, Shuqi LI, and Gadeng Nima on June
4, 2017. KIZ 044431, adult male, KIZ 044432, 044433, adult fe-
males, from Rongzong Village, Deqin County, Yunnan Province,
China (99.1733° E, 28.5804° N, elevation 2,198 m, WGS 84); KIZ
044740, adult male, KIZ 0 44742, adult female, near Zhidu, Deqin
County, Yunnan Province, China (99.2241° E, 28.3545° N, elevation
2037 m, WGS 84); CIB 116315–16, subadult females, CIB 116318,
adult male, all from Songmai Township, Derong County, Sichuan
Province (99.292382° E, 28.701581° N, elevation 2520m, WGS 84),
collected by Gang Wang and Puyang Zheng on August 29, 2019; KIZ
044764, adult male near Derong township, Sichuan Province, China
(99.2773° E, 28.6714° N, elevation 2,412 m, WGS 84); all collected
by Kai WANG, Zhuoyu LU, and Xiankun HUANG on June 19, 2019.
Diagnosis
The new species can be diagnosed from congeners by a combination
of the fol low in g mor pho log ica l cha rac ter ist ics : (1) bo dy size mod era te,
SVL 56.3–61.2 mm in males, 57.0–66.5 mm in females; (2) tail long,
TAL 212.0%–221.1% SVL in males, 191.4%–207.7% in females; (3)
hind limbs moderate, HLL 73.7%–82.7% SVL in males, 71.8%–80.5%
in females; (4) head moderate, HW 66.8%–75.1% HL; (5) MD 36–45;
(6) T4S 19–24; (7) post-rictal conical or sub-pyramidal scales weak and
few, 1–3; (8) tympanum concealed; (9) nuchal crest well-developed on
strong skin folds, serrated; (10) distinct transverse gular fold present;
(11) ventral scales of head and body distinctively keeled; (12) ven-
tral head scales and ventrolateral body scales homogeneous in size;
(13) gular spots present in both sexes, Pale Cyan (Color 157) to Light
Caribbean Blue (Color 163) in life, Plumbeous (Color 295) after long-
term preservation; (14) dorsolateral stripes strongly jagged, Cream
(Color 12) in males, white in females in life; (15) dark ornamentations
(i.e., transverse bands, radial stripes around eyes) with strong con-
trast; (16) no distinct ornamentation pattern on ventrolateral body;
(17) ventral body white in most individuals, sometimes pale yellowish
in males; and (18) oral cavit y, inner lips, and tongue light flesh color
(Color 250).
Comparisons
The new species was confused with D. flaviceps, but it can be differ-
entiated from the latter by having a smaller body size in males (SVL
56.3–61.2 mm versus 68.5–82.1 mm), fewer middorsal crest scales
(MD 35–46, average 40 versus 45–56, average 51), solid patches
along dorsal midline in both sexes (versus hollow, rhomboid-shaped
patterns), as well as by the presence of distinct radial stripes around
eyes (versus absence or faint), the presence of gular spots in both
sexes (versus absence), the absence of crest skin folds in females
(versus absence), and the absence of reticulated, vermiculate pat-
terns on the gular region (versus presence).
Diploderma aorun sp. nov. is morphologically most similar to
D. batangense, which is found further upstream along the Jinsha
River. However, D. aorun sp. nov. can still be differentiated from
D. batangense by having a longer tail in males (TAL 212.0%–221.1%
SVL, average 216.4% versus 187.3%–206.5%, average 194.6%;
Tukey's HSD test, p < .05), strongly developed skin fold of nuchal
and dorsal crests in males (versus weak and often indistinct), and
much stronger contrast between the dorsal dark and light ornamen-
tations (versus much weaker contrast and faint).
FIGURE 9 Male holotype (KIZ 0 40138) of Diploderma panlong sp. nov. in life. Photographs by KW
(a) (b) (c)
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239
WANG et Al.
For the remaining species in the same clade (Figure 2), Diploderma
aorun sp. nov. differs from D. drukdaypo, D. vela, and D. slowinskii by
the presence of distinct gular spots in both sexes (versus absence in
both sexes); from D. iadinum b y hav ing fewe r dor sol ate r al rows of en-
larged keeled scales on each side of the body (1 versus 2–3), strongly
jagged dorsolateral stripes (versus smooth), distinct body coloration
in both sexes (Pale Buf f [Color 1] in males, Chamois [Color 84] in
females versus Yellowish Spectrum Green [Color 128] to Emerald
Green [Color 143] in males, Buff [Color 5] to Pale Greenish Yellow
[Color 86]), and a distinct gular coloration (Pale Cyan [Color 157] to
Light Caribbean Blue [Color 163] in both sexes versus Caribbean Blue
[Color 168] in males, Medium Greenish Yellow [Color 88] in females);
from D. yulongense by having a distinct coloration for gular spots in
life (Pale Cyan [Color 157] to Light Caribbean Blue [Color 163] ver-
sus Chartreuse [Color 89] to Opaline Green [Color 106]) and dorso-
lateral stripes (white to Cream Color [Color 12] versus Pale Greenish
Yellow [Color 86]), as well as by the absence of green patches on the
dorsolateral surface of body (versus presence); from D. laeviventre by
having distinctively keeled ventral scales (versus smooth or feebly
keeled) and distinct gular coloration (Pale Cyan [Color 157] to Light
Caribbean Blue [Color 163] versus Light Chrome Orange [Color 76]);
from D. chapaense and D. yunnanense by the presence of a distinct
transverse gular fold (versus absence) and differential coloration of
gular spot (Pale Cyan [Color 157] to Light Caribbean Blue [Color 163]
versus Light Chrome Orange [Color 76]).
For the remaining species in the different phylogenetic clade,
D. aorun sp. nov. differs from D. micangshanense, D. varcoae, an d
all species from East Asian islands (D. brevipes, D. luei, D. makii,
D. polygonatum, and D. swinhonis) by the presence of a distinct
transverse gular fold (versus absence); from D. dymondi, D. swild,
D. slowinskii, and D. varcoae by having a concealed tympanum
(versus exposed) and distinct coloration of oral cavity (Light
Flesh Color [Color 250] versus Light Chrome Orange [Color 76] in
D. swild and D. varcoae; Jet Black [Color 300] or Spectrum Violet
[Color 186] in D. dymondi); from D. splendidum by having distinc t
transverse gular fold (versus feeble), homogeneous ventral head
scales (versus heterogeneous), and jagged dorsolateral stripes in
males (versus smooth-edged); and from D. zhaoermii by having a
smaller body size (maximum SVL 61.2 mm in males versus up to
81.7 mm), distinc t vermiculate stripes on the ventral head (versus
faint or absence), and a distinct gular coloration (Pale Cyan [Color
157] to Light Caribbea n Blue [Color 163] versu s Ch ar treu se [Color
89]).
For the species that lack genetic data, D. aorun sp. nov. differs
from D. brevicaudum by having a longer tail (TAL ≥ 191.4% versus
≤160.0%) and longer hind limbs (>71.5% versus ≤64.0%) and the
presence of gular spots after preservation (versus absence); from
D. grahami by having distinct appearance of dorsal body scales (spiky
versus granular), a distinc t transverse gular fold (versus feebly), and
a larger body size (56.3–66.5 mm versus 49.3 mm); from D. fasciatum
by having smaller and non-dif ferentiated nuchal crest scales (ver-
sus much larger and distinctively differentiated from dorsal crests),
the presence of dorsolateral stripes (versus absence), and by the ab-
sence of single hourglass-shaped pattern on the mid-dorsum (ver-
sus presence); and from D. hamptoni by having parallel dorsolateral
stripes on dorsal body (versus diagonally away from dorsal midline).
For new species described above, D. aorun sp. nov. differs from
D. angustelinea sp. nov. by having wider and strongly jagged dorso-
lateral stripes in males (versus much narrower, smooth or feebly jag-
ged), a distinct gular spot coloration (Pale Cyan [Color 157] to Light
Caribbean Blue [Color 163] versus Dark Spectrum Yellow [Color
78]), a muc h gradual tra nsition from dorsal to ventr al coloration (ver-
sus sharp and distinct), and by the presence of strongly developed
skin folds under nuchal and dorsal crests in males (versus absence),
FIGURE 10 Male holotype (a1–a3; KIZ 028332) and female paratype (b1–b3; KIZ 044820) of Diploderma qilin sp. nov. in life. Photographs
by KW
(a1) (a2) (a3)
(b1) (b2) (b3)
240
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WANG et Al .
and the presence of distinct black vermiculate stripes on ventral
head (versus absent).
Description of holotype
Body size moderate, not dorsally compressed, SVL 56.3 mm; tail
slender, long, TAL 213.63% SVL; limbs moderate, FLL 48.3% SVL,
HLL 80.8% SVL; head relatively wide and robust, HW 72.7% HL, HD
75.8% HW; snout pointy, SEL 40.8% HL. Rostral rectangular, three
times wider than height, two scales away from nasal; nasal oval, sin-
gle scale away from first supralabial; supralabials 9/8, feebly keeled,
more dis ti nct on la st fi ve; infr al abials 11/11 , feebl y ke el ed; subor bi tal
scale rows 3/3, middle row slightly enlarged, each scale bearing dis-
tinct, lateral keel; supraciliaries 6/6, overlap one half of total length
with succeeding ones; enlarged, protruding scales between posterior
orbit and tympanum, 6/7, each bearing single distinct, lateral keel;
tympanum concealed under fine scales; enlarged, conical scales
post-tympanic, 6/5, one of which much taller than others on each
side; enlarged post-rictal scales 4/5, sub-pyramidal in shape. Dorsal
head scales heterogeneous in size and shape, all distinctively keeled;
single, laterally oriented, Y-shaped ridge present on dorsal snout,
starting three scales posterior of rostral to posterior end of orbit;
interparietal slightly enlarged, parietal eye distinct; postciliary scale
much enlarged, subpyramidal shape, with multiple keels; post-occip-
ital scales well-differentiated, conical or subpyramidal in shape, 4/3.
Dorsal body scales distinctively keeled, heterogeneous in size
and shape; axillary scales fine, much smaller than remaining dor-
sals; enlarged scales mostly randomly scattered, except ones close
to dorsal midline and ones along medial line of dorsolateral stripes,
which arranged in dorsolateral rows from neck to pelvis on each
side; nuchal, dorsal crests well-developed on skin folds, crest scales
erected, serrated, differentiated from nearby dorsal sales; middorsal
scales 38. Scales of dorsal limbs distinc tively keeled, homogeneous
on forelimbs, heterogeneous on hind limbs, with few enlarged, sub-
pyramidal ones on posterior lateral thigh and dorsal crus; Finger IV
and Toe IV longest, Finger IV subdigital lamellae 14/15, Toe IV sub-
digital lamellae 21/21.
Vent ral he ad scales homoge ne ou s in size and sha pe , dist in ct iv el y
keeled except chin shields, regularly arranged; mental pentagonal,
not completely enclosed by first pair of chin shields; chin shields
5/5, smooth, two scale rows from infralabials. Gular pouch present,
well-developed in life, indistinct after preservation; distinct trans-
verse gular fold present, deep. Ventral body and limb scales distinc-
tively keeled, keels more pronounced than those of ventral head,
homogeneous in size and shape, mostly regularly arranged.
Coloration of holotype in life
The background coloration of the dorsal surface of the head is
Smoky White (Color 261). Four Dark Neutral Gray (Color 299)
transverse bands are evenly distributed on the dorsal surface of
head, with the last one between posterior margins of the orbits. A
thin, incomplete, transverse band is between the 2nd and the 3rd
and be tween 3rd and 4t h thick ban ds . A ll of these transver se band s
except the most anterior three enter orbits on both sides, which
form the radial stripes superior of the eyes. Temporal and occipit al
regions of the head are dirty Light Neutral Gray (Color 297) to Jet
Black (Color 300), with irregular Dark Neutral Gray (Color 299) to
Jet Black (Color 300) spots and streaks. The ground coloration of
the lateral surface of the head is white. Jet Black (Color 30 0) radial
stripes are present around eyes, where the posterior two stripes
are the broa de s t: On e of wh ich exte nds from the su per io r pos ter ior
corner of orb it to region above the concealed t ympanum, while th e
other extends from inferior posterior corner of orbit to region just
su per ior to the ri ctu s. Som e Irr egu lar Dark Ne utr a l Gra y (Co lor 299)
speckles are present on the lateral surfaces of head, particularly
posterior to orbits.
The ground coloration of the dorsal and lateral surface of body
is Pale Buff (Color 1). A single strongly jagged, Cream Colored (Color
12) dorsolateral stripe is present from neck to pelvis on each side
of the body. Six Jet Black (Color 30 0) rectangular patches are pres-
ent on the dorsal body along dorsal midline from neck to pelvis
betw ee n dorso la te ra l str ip es. Eac h of the rect angular patches is se p-
arated from each other by s Cream Colored (Color 12) transverse
streak, which connects the dorsolateral stripes on both sides of the
body. Lateral surfaces of the body that are inferior to dorsolateral
stripes are Jet Black (Color 300), with Cream Colored (Color 12)
scales densely scattered. Dorsal sur faces of limbs are white, with
Dark Neutral Gray (Color 299) to Jet Black (Color 300) transverse
bands across. The bands are much distinct on forelimbs than hind
limbs. The dorsal surface of tail is Smoky White (Color 261). Faint
Dark Neutral Gray (Color 299) or Jet Black (Color 30 0) transverse
bands are evenly scattered on the tail from the vent to two third of
its length, and the remaining distal portion of the tail is more or less
uniform Drab-Gray (Color 256).
The ground coloration of the ventral surface of the head is
white. Short, Jet Black (Color 300) streaks and speckles are pres-
ent, some of which connect and form vermiculated patterns. A dis-
tinct Medium Blue (Color 169) gular spot is present on the posterior
center of ventral head, relatively large in size. Such gular spot ends
before the transverse gular fold and does not extend into anterior
chest. The ventral surface of the body is uniform white without any
distinc t ornamentation. The hands and feet are uniform pale Tawny
Olive (Color 17), while the remaining ventral sur faces of limbs are
uniform white. The ventral surface of the anterior two third of tail
is white, while the remaining parts gradually become Smoky White
(Color 261) toward the tip.
Coloration of holotype in preservation
Ornamentation remains largely the same after preservation, but
coloration fades. Specifically, the Cream Color (Color 12) of dor-
solateral stripes and scattered scales on the lateral surface of the
body fade into white, and the Medium Blue (Color 169) gular spot
becomes Light Sky Blue (Color 191).
Variatio n
Morphometric and pholidosis variation of the type series is sum-
marized in Table S8. The dorsal coloration of a single male (KIZ
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WANG et Al.
044740) is Vandyke Brown (Color 282) to Jet Black (Color 300),
with no distinct dorsolateral stripes. For the coloration of the gular
spot, some individuals (i.e., KIZ 032734, 032736, 032737) are
much lighter than others, showing a Pale Cyan Color (Color 157).
Although sample size is limited, sexual dimorphism is evident in
the species, where similar to congeners, males of D. aorun sp. nov.
show tendency toward a longer tail (TAL 212.0%–221.1% SVL in
males versus 178.85%–207.7% in females). Furthermore, female
specimens possess distinct coloration and ornamentation patterns
compare to males, including having a distinct coloration of dor-
solateral stripes (Smoky White [Color 261], Sulphur Yellow [Color
80], or entirely absent in females versus Cream Color [Color 12] in
males), background coloration of lateral head, lateral body inferior
to dorsolateral stripes, and dorsal limbs (Chamois [Color 84] in live
females versus white in males), and by the presence of Peach Red
(Color 70) vertebral stripe pattern from dorsal head to the vent in
female (versus absence in males). Males tend to have brighter and
larger gular spots than females.
Etymology
The species name “aorun” is derived from the name of the legendary
Dragon Lord in Chinese mytholog y (Chinese 敖闰, Pi nyi n: Ao Run), who
is the guardian of the West Ocean and is responsible for creating pre-
cipitation. We name the new species after the guardian of the ocean
in the hope that the recognition of its endangered status will bring
protection to the fragile valley habitats that the species is endemic
to, just like the dragon lord protec ting its realm in the mythology. We
recomm end Aor un Mountain Dr agon as its Engli sh co mmon name an d
敖闰龙蜥 (Pinyin: Ao Run Long Xi) as its Chinese common name.
Natural Histor y and conservation
Diploderma aorun sp. nov. inhabits hot–dry valley below 3,000 m
of elevation along the upper Jinsha River, from Zongrong Village
of Derong County, Sichuan Province, to areas 14km south of
Benzilan, Deqin County, Northwest Yunnan Province, China. Its
habitat is the most arid among all new species described here,
which comprises of most rock y outcrops (mostly sandstone) and
very little vegetation (Figure 13b). The species is terrestrial, bask-
ing on large rocks or open grounds within patches of cactus (an
introduced alien species) during the day, but most individuals were
observed sl ee ping on bushes or gra ss stems at nig ht . Roa d-ki ll indi-
vid ua ls were commonly seen along th e highway thr ou gh the val ley.
Elaphe taeniura and Gekko scabridus were found co-distributed at
the type locality.
FIGURE 11 Comparisons of lateral head view (1), dorsal head view (2), ventral head view (3), dorsolateral overview (4), and ventral
overview (5) of preser ved specimens among males of Diploderma angustelinea sp. nov. (a), D. aorun sp. nov. (b), D. flavilabre sp. nov. (c), D. qilin
sp. nov. (d), and D. panlong sp. nov. (e). Photographs by KW
(a1) (a2) (a3) (a4) (a5)
(b1) (b2) (b3) (b4) (b5)
(c1) (c2) (c3) (c4) (c5)
(d1) (d2) (d3) (d4) (d5)
(e1) (e2) (e3) (e4) (e5)
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Based on our sur veys, the known extent of occurrence of
D. aorun sp. nov. is only approximately 550 km2, which includes
low-elevation (i.e., <3,500 m), hot–dry valleys along the isolated
upper Jinsha River (from Benzilan to Zongrong Village) and its im-
mediate tributaries (i.e., Dingqu and East Wangxuqu). Although
the Baima Snow Mountain Nature Reserve protects the west side
of the Jinsha Valley within the range of D. aorun sp. nov., being
the most peripheral part s of the reserve, much of the habitats still
overlap greatly with rapid human developments, national high-
ways, and tourist infrastructures. In fact, multiple illegal cement
mills were observed at the type locality of the species, which
devastate the limestone habitats around them. Additionally, the
recent major landslides in 2018 and the discharge of its resulting
barrier lake on the Jinsha River have led to severe habitat destruc-
tions along the Jinsha Valley, and the follow-up road repairs after
the natural disaster further destroyed the low-elevation habitats.
Despite the abundant individuals in some regions, the population
density varies drastically across its range, and numerous human
settlements fragment the continuous distribution of the species,
leading to isolated populations along the valleys. Therefore, fol-
lowing IUCN criteria D2, we recommend listing D. aorun sp. no v.
as Vulnerable (VU).
3.3.3 | Diploderma flavilabre sp. nov. Wang,
Che, Siler
(Population 12 in Figure 1; Figures 2, 7, 11, 12;
Tables 2–4; Table S8 (3); ZooBank ID urn:lsid:zoobank.
org:act :90 059942-6615-4C6A-834A-C427B1A A9BE6).
Synonym
Japalura flaviceps Zhao et al. (1999: 111–115), in part; Zhao (2003:
84), in part.
Holotype
KIZ 032693, adult male from Jinsha River Valley at Yebatan, Gaiyu
County, nor thwest Sichuan Province, China, on May 28, 2017
(98.9642° E, 30.7385° N, elevation 2,845 m, WGS 84).
Paratopotypes
KIZ 032692, 032694, adult males; KIZ 032695–032699, KIZ 032730,
adult females; all collected from the same locality as the holotype by
Kai WANG and Gadeng NIMA.
Diagnosis
The new species can be diagnosed from congeners by a combination
of the following morphological characteristic s: (1) body size small,
SVL 50.8–55.2 mm in males, 64.2–72.9 mm in females; (2) tail short,
TAL 149.7%–179.3% SVL in males, 140.2%–152.2% in females; (3)
head wide, HW 70.01%–73.0% HL, HD 68.6%–75.5% HW; (4) hind
limb short, HLL 66.9%–74.8% SVL in males, 61.7%–63.9% in females;
(5) tympanum concealed; (6) transverse gular fold present, deep;
(7) MD 39–43; (8) F4S 13–16; (9) T4S 19–22; (10) post-rictal conical
scales well-developed, 4–9; (11) nuchal and dorsal crest well-devel-
oped with strong skin folds in males only; (12) ventral head scales
smo oth or fe eb ly keeled; (13) vent ra l body scales dis tinc ti vely keeled;
(14) inner lips Dark Spectrum Yellow (Color 78) in both sexes in life,
distinc t from background coloration of oral cavity; (15) background
coloration of oral cavity Light Flesh Color (Color 250) in life; (16) pal-
ate and tonsils speckled heavily with Light Neutral Gray (Color 297) in
both sexes in life; (17) gular spots present in both sexes, Pale Emerald
Gr een (Co lor 141) to Ligh t Tur quo is e Gre en (C ol o r 146) in life ; an d (18)
ventral body Pale Greenish Yellow (Color 86) in live males.
Comparisons
The new species was confused with Diploderma flavicpes, but it can
be distinguished from D. flaviceps readily by having a much smaller
adult body size in males (SVL 50.8–55.2 mm versus 68.5–82.1 mm),
a relatively shorter tail (TAL 149.7%–179.3% SVL in males, 140.2%–
152.2% in females versus 181.8%–210.8% in males, 191.5%–205.7%
in females), distinct coloration of inner lips (dark spectrum yellow
coloration versus same as rest of oral cavity, flesh color), as well as by
the presence of gular spots in both sexes (versus absence), presence
of distinct radial stripes around eyes (versus faint or absence), pres-
ence of distinct dark spectrum yellow along inner lips in both sexes
(versus absence), absence of reticulated dark patterns on gular re-
gion (versus presence), and by the absence of hallow, brown rhomb-
shaped patterns with distinct yellow centers along dorsal midline
(versus presence).
Diploderma flavilabre sp. nov. is morphologically most similar
to D. batangense and D. vela, with all three species possessing dis-
tinct black radial stripes around eyes, blackish dorsal background
coloration, and raised nuchal and dorsal crests on skin folds in
males. However, the new species can be differentiated from both
D. batangense and D. vela by having a distinct coloration color of
inner lips in life for both sexes (Dark Spec trum Yellow [Color 78]
versus flesh color, same as the rest of oral cavity) and the ten-
dency toward a relatively shorter t ail (ANOVA p = .031; TAL
149.7%–179.3% SVL in males, 140.2%–152.2% in females versus
187.3%–206.5% in males, 171.8%–203.1% in females for D. batan-
gense [Tukey's HSD p = .041]; 174.6%–238.1% in males, 159.8%–
202.7% in females for D. vela [Tukey's HSD p = .034]). In addition,
it differs from D. batangense by having more post-rictal modified
conical or sub-pyramidal scales (4–9 versus 0–3); and from D. vela
by having a relatively wider head (HW 70.1%–73.0% HL versus
65.2%–69.8%), discontinuous skin folds between nuchal and dor-
sal sections in males (versus continuous), distinct coloration of
dorsolateral stripes in females (Sulphur Yellow [Color 80] versus
Medium Chrome Orange [Color 75]), and by the presence of gular
spots in both sexes (versus absence).
For remaining closely distributed congeners in the northern
HMR, the new species can be differentiated from all species by
the distinct dark spectrum yellow coloration of inner lips in life
for both sexes (versus same flesh color as the rest of oral cavity).
Furthermore, it can be dif ferentiated from D. brevicaudum by the
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WANG et Al.
presence of distinct gular spots in both sexes after preservation
(versus absence); from D. laeviventre and D. drunkdaypo by having
dist inc t iv ely kee led vent ral head and bo dy sca les (ve rsu s smo ot h or
feebly keeled); from D. slowinskii by a smaller body size (SVL 50.8–
55.2 mm in males, 64. 2–72.9 mm in females versus 81.0–95.5 mm
in males, 74.8–91.7 mm in females) and a concealed tympanum
(versus exposed); from D. yulongense by having a much shorter
tail (TAL 149.7%–179.3% SVL in males, 140.2%–152.2% in females
versus 211.3%–247.6% in males, 182.1%–227.3% in females), as
well as by the absence of greenish ventrolateral patches of body
(versus presence); and from D. iadinum by the distinct body col-
oration in males (Jet Black [Color 300] background coloration of
dorsal body, with Pale Horn Color [Color 11] dorsolateral stripes
versus Yellowish Spectrum Green [Color 128] to Emerald Green
[Color 143] background coloration, with same colored dorsolat-
eral stripes), shape of dorsolateral stripes in males (strongly jagged
versus smooth-edged), and fewer dorsolateral ridges on body in
females (1 versus 2 or 3).
For the remaining recognized congeners, in addition to the dif-
ferential Dark Spectrum Yellow (Color 78) coloration of inner lips
(versus same coloration as in the oral cavit y), the new species dif-
fers from D. varcoae, D. dymondi, and D. swild, by having a concealed
tympanum (versus exposed); from D. chapaense, D. grahami, D. mi-
cangshanense, D. yunnanense, and all island species (D. brevipes,
D. luei, D. makii, D. polygonatum, and D. swinhonis) by having a distinct
transverse gular fold (versus absence or feeble); from D. fasciatum by
the absence of single hourglass-shaped pattern on the mid-dorsum
(versus presence); from D. splendidum by much smaller adult body
size (SVL 50.8–55.2 mm in males, 64.2–72.9 mm in females versus
>80.0 mm in both sexes) and much shorter tail ( TAL < 180.0% SVL
in males, <153.0% in females versus >200% in both sexes); and from
D. zhaoermii by a smaller adult body size in males (SVL < 55.2 mm
versus > 64 mm) and more T4S (T4S 19–22 versus 23–27).
Lastly, for the other new species described here, D. flavilabre
sp. nov. differs from all by the dif ferential coloration of inner lips
(versus same coloration as rest of the oral cavity). Specifically, it
differs from D. angustelinea sp. nov. by having a relatively shorter
tail (TAL 149.7%–179.3% SVL in males, 140.2%–152.2% in females
versus 230.2%–249.1% in males, 194.3%–222.3% in females), more
and better developed conical or sub-pyramidal post-rictal scales
(4–9 versus 0–3), distinct coloration of gular spots (Pale Emerald
Green [Color 141] versus Spectrum Yellow [Color 79]), wider and
strongly jagged dorsolateral stripes (versus narrow, weakly jagged
or smooth), as well as by the presence of raised, well-developed nu-
chal and dorsal crests on skin folds in males (versus absence); and
from D. aorun sp. nov. by having a smaller body size in males (SVL
50.8–55.2 mm ver sus 58.4–61.2), a shorter t ail (TAL 149.7%–179.3%
SVL in males, 140. 2%–152.2% in females versus 212.0%–221.1% in
males, 191.4%–207.7% in females), shorter hind limbs (HLL 66.9%–
74.8% in males, 62.0%–65.6% in females versus 73.7%–82.7% in
males, 71.8%–80.5% in females), and a distinct gular coloration (Pale
Emerald Green [Color 141] versus Pale Cyan [Color 157] to Light
Caribbean Blue [Color 163]).
Description of holotype
Small-sized agamid, SVL 50.8 mm; tail short, TAL 172.8% SVL;
limbs relatively short, FLL 48.3% SVL , HLL 74.8% SVL; head
wide, robust, HW 71.2% HL, HD 75.5% HW; snout pointy, SEL
37.1% HL. Rostral rectangular; nasal oval-shaped, nostril close to
posterior end, two scales away from the rostral, one scale away
from first supralabial; supralabials 9/8 (left/right), feebly keeled;
infralabials 10/8, smooth; loreal approximately equal size, irregu-
larly arranged, keeled; suborbital scale rows 3/4, keeled, central
row slightly enlarged; supraciliaries 7/7, each greatly overlapped
two third total length with posterior scale. Tympanum concealed
under fine scales; two enlarged, sub-pyramidal scales anterior
tympanum; enlarged, conical, post-tympanic scales 6/6, one of
which distinctively taller on each side; enlarged, conical or sub-
pyramidal, post-rictal scales 8/8, one/two much taller than others
on eac h si de. D or sal h ea d sca le s ke eled, het er og en eous in size an d
shapes; enlarged, hexagon-shaped scales forming Y-shaped ridge
on do rs al snout anter ior to eyes; interp ariet al not much dif ferenti-
ated, parietal eye indistinct; single enlarged, sub-pyramidal scale
dorsoposterior to orbit on each side of head; enlarged conical
scales 7/5 post-occipital.
Dorsal body scales heterogeneous, strongly keeled; axillary
scales granular, much smaller than other dorsal scales; enlarged,
sub-pyramidal scales, each with single distinct medial keel, scat-
tered across dorsum; enlarged scales mostly irregularly arranged,
except some arranged in paravertebral row superior to dorsolat-
eral stripes on each side of body. Middorsal scales 41; nuchal crest
scales serrated, erected, much larger than neighboring dorsal
scales; dorsal crest scales less erected than nuchal, serrated; both
cres ts on well-develo ped skin folds in life, with distinct bre ak in be-
twee n; bot h skin fold s of cre sts shrunken aft er preser vation. Do rsal
limb scales distinctively keeled, homogeneous in size on forelimbs,
heterogeneous on hind limbs, with some enlarged sub-pyramidal
ones on posterior lateral thigh and dorsal crus. Finger IV and Toe
IV longest, Finger IV subdigital lamellae 16/16, Toe IV subdigital
lamellae 20/20.
Ventral head scales homogeneous in size and shape mostly, with
few slightly enlarged toward center of gular, regularly arranged,
smooth or feebly keeled; mental pentagonal in shape, enclosed by
first pair of chin shields, which compressed and elongated; chin
shield 5/4, smooth, 2–3 scale rows from infralabials. Gular pouch
present in life, well-developed; distinct transverse gular fold present
across neck, deep. Ventral body scales mostly homogeneous in size
and shape, distinctively keeled. Ventral limb and tail scales distinc-
tively keeled; ventral thigh scales regularly arranged in rows in proxi-
mal–distal direction, smaller toward posterior lateral sides.
Coloration of holotype in life
The background of the anterior part of dorsal head is white to Smoky
White (Color 261). A single V-shaped, Jet Black (Color 300) pat-
terns on the dorsal anterior snout, with it s branches terminate on
the nasal scales, and the root terminates at the posterior edge of
the first enlarged, keeled scale of the Y-shaped ridge on the snout.
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Three Orange-Rufous (Color 56) transverse bands with distinct Dark
Neutral Gray (Color 299) edges are present on the dorsal surface of
head, where the first one is short and centered on the dorsal snout
between the anterior borders of orbit cir cle. The backg ro un d co lo ra -
tion of the temporal region of the head is Smoky Gray (Color 266),
with irregular ornamentation patterns of Ducky Brown (Color 285)
to Jet Black (Color 30 0).
Lateral surface of head is white to Light Buff (Color 2). Distinct
Jet Black (Color 300) radial stripes are around the eyes, eight and
nine on each side. Of these radial stripes, three stripes are below
eyes in subocular and loreal regions, and all of them continue to ex-
ten d to lower jaw. The two pos te ri or ly dir ec ted strip es fro m th e po s-
terior corner of each eye (one extending slightly dorsally, the other
extending slightly ventrally) are the broadest among all, where the
ventrally extending one terminated at the anterior end of tympanic
region. The inner surfaces of both upper and lower lips, as well as
corners of the mouth, are Dark Spectrum Yellow (Color 78). The
tongue and gum are uniform Light Flesh Color (Color 250), whereas
the palate and tonsils are speckled heavily with Light Neutral Gray
(Color 297).
Background coloration of the dorsal surface of the body is
Jet Black (Color 300). A distinct Pale Horn Color (Color 11), zig-
zag-shaped, dorsolateral stripe is present on each side of the ver-
tebral crest, running from neck to pelvis, sympatric to each other
along the vertebral line. The point y edges of the jagged dorsolat-
era l stri pe extend dorsall y an d almost conn ec t wi th t he sym metric
edges from the other side at the dorsal midline. A s the individual
become stressed or when it was basking, the coloration of dor-
solateral stripes may become dirty with pale Salmon Color (Color
58). A total of six solid, dark patches are present along the dor-
sal midline between two dorsolateral stripes from neck to vent,
which are Mahogany Red (Color 34) in the center and gradually
transition to Jet Black (Color 300). Except the first and the fourth
patches, which are in rectangular and irregular shape, respectively,
the remaining four dark patches are in rhomb shape. Irregular light
spots of white to Pale Pinkish Buff (Color 3) coloration are scat-
tered inferior to the dorsolateral stripe on each side. Those light
spots become larger and longer as moving toward ventral direc-
tion, which separates the background Jet Black (Color 300) color-
ation into thin reticulated patterns on ventrolateral sides of body.
Dorsal surfaces of limbs are Jet Black (Color 30 0). Distinct Pale
Buff (Color 1) to Pale Pinkish Buff (Color 3) transverse bands are
present on dorsal limbs, evenly scattered from proximal to distal
end. Similar transverse bands are also present on dorsal surfaces
of limb digits. A white, narrow stripe is present from the base of
thigh to the knee joint on the posterior ventrolateral side of hind
limb on each side.
The background coloration of the ventral sur face of the head
is white. A relatively large, two-centered arch-shaped, gular spot
is present on the posterior central region of the ventral head, Pale
Emerald Green (Color 141) in coloration. Jet Black (Color 300) ver-
miculated stripes are present on the remaining areas of ventral
head out side of the gular spot . Vent ral sur face of neck and the very
anterior portion of the ventral surface of the chest are also Pale
Emerald Green (Color 141) as in the gular spot. This Pale Emerald
Green coloration gradually fades into Pale Greenish Yellow (Color
86) as moving pos teriorl y to the chest and abdo men; an d colorat ion
continues to fade, where it eventually becomes Pale Buff (Color 1)
at the very posterior portion of the ventral body. No other distinct
ornamentation patterns are present on ventral surface of body.
Ventral surfaces of limbs are uniform Pale Buff (Color 1), which
becomes darker and transitions to Pale Pinkish Buff (Color 3) on
ventral hands and feet.
Background coloration of the tail is Pale Pinkish Buff (Color 3).
Distin ct , da rk transvers e bands are pres ent on dorsal an d lat eral sur-
faces of t ail, which do not fully enclose the tail on the ventral side.
Coloration of transverse bands changes from Medium Neutral Gray
(Color 298) at the proximal end to Salmon Color (Color 58) at the
distal end.
Coloration of holotype in preservation
While the ornamentation patterns remain the same after preserva-
tion, coloration of the holotype fades significantly. Specifically, the
Pale Horn (Color 11) coloration of the dorsolateral stripes, the Pale
Pinkish Buff (Color 3) coloration of irregular spots on the dorsal
surface of the body, and the Dark Spectrum Yellow (Color 78) col-
oration of the inner lips fade into white; the Mahogany Red (Color
34) coloration of the dorsal body becomes Burnt Umber (Color 48);
the Pale Emerald Green (Color 141) gular spot fades into Pale Cyan
(Color 157); and the Pale Greenish Yellow (Color 86) coloration of
the anteroventral region of the body fades into Cyan White (Color
156) or white.
Variatio n
Morphometric and pholidosis variation of the type series is summa-
rized in Table S8. Although the sample size is small, it is evident that
the new sp ecies is sexually dimorp hic. Ma les pos sess relat ive ly a rel-
ativel y lo nger ta il (TA L 172 . 8%–179.3% SV L in male s ve rsu s 140. 2%–
152.2% in females), relatively longer hind limbs (HLL 66.9%–74.8%
SVL in males versus 61.7%–65.6% in females) and raised crests on
skin folds (versus absence). Furthermore, males of the new species
differ from females in coloration and ornamentation patterns, in-
cluding having distinct background coloration of dorsum (Jet Black
[Color 300] to Mahogany Red [Color 34] in males versus Tawny Olive
[Color 17] to Clay Color [Color 18] in females) and presence of dis-
tinct Pale Horn Color (Color 11) dorsolateral stripes in males (versus
absence or in faint Sulphur Yellow [Color 80]).
Among females, the transverse bands on the dorsal head and
dark rhomb-shaped patches along dorsal body midline of KIZ
032697 are Light Pratt's Rufous (Color 71); the same specimen
also has irregular Drab-Gray (Color 256) patterns on the ante-
rior and ce nt ral a bd omen, and it s gula r spot is Gr ee nish Turqu oi se
(Color 166). KIZ 032699 has the faintest vermiculated stripes on
the ventral head. KIZ 032700 has Grayish Horn (Color 268) color-
ation, irregular stripes and patches on the chest region of ventral
body.
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Etymology
The Latin species name, flavilabre, means “yellow-lipped,” which de-
scribes the diagnostic dark spectrum yellow coloration of inner lips
of the species. We recommend Yellow-lipped Mountain Dragon as
its English common name and 黄唇龙蜥 (Pi nyin: Hua ng Chun Long Xi )
as its Chinese common name.
Natural Histor y and Conservation
The overall habitat of the new species is warm, semi-arid valley
with considerable amount of vegetation coverage, which is ver y
distinc t from habitats of congeners in lower reaches of the river
(e.g., D. batangense and D. aorun sp. nov.). Individuals were observed
inhabiting rock piles (sandstone) among shrubs and forest edges
(Figure 13c). Females and juveniles seem to be less selective for
weather conditions and were active even in during cloudy weather,
whereas males were only observed during the hottest time of the
sunny days. All observed females, including ones that were col-
lected, were gravid, suggesting June is during the breeding season of
the species. Considerable amount of ant remains were found in the
feces of the collected individuals. Possible predations are likely from
birds and snakes (Elaphe taeniura).
The type locality of D. flavilabre sp. nov. is less than five km
away from the biggest hydropower station in the upper Jinsha
River, the Yebatan Hydropower Station, which was under active
construction at the time of collection in 2017. Several habitat de-
structions were observed at the time, with explosions and drain-
age alterations. No lizards were found close to the construction
site, and population density was much lower than any other new
species described here. Furthermore, the recent major land-
slide and subsequent flooding in 2018 might have led to further
massive habitat destructions and population decline of the new
species.
Because we did not find any D. flavilabre sp. nov. near Baiyu
Township, which is approximately 60 km linear distance north of the
type locality of D. flavilabre sp. nov., the potential distribution range
of the new sp eci es is only less than 140 k m lin ear dis t anc e of low- el e -
vation habitats along a single, isolated river valley, and the estimated
extent of occurrence is less than 400 km2. Given the endemic na-
ture and limited range, fragile ecosystem where the species inhabits,
and several, continuous habitat destructions from both natural and
anthropogenic causes, we propose to list the species as Vulnerable
(VU) based on IUCN criteria D2.
3.3.4 | Diploderma panchi sp. nov. Wang, Zheng, Xie,
Che, Siler
(Population 6 in Figure 1; Figures 2, 8, 12; Tables 2–4;
Table S8 (4); ZooBank ID urn:lsid:zoobank.org:act:833577B9-
CF79-4E8E-AC83-D8E913DF8DBA).
Synonym
Japalura flaviceps Zhao et al. (1999: 111–115); Zhao (2003: 84).
Holotype
KIZ 032715, adult female from upper Yalong River Valley near Yajiang
Township, Yajiang County, Ganzi Tibetan Autonomous Prefecture,
northwe st Sichua n Prov ince, Chi na (30. 0 414° N, 101. 0108 ° E, el ev a-
tion 2,663 m). Collected by Kai WANG and Nima DADENG in May
22, 2017.
Paratopotypes
KIZ 032716, 032717, 032729, adult females, same collecting infor-
mation as for the holot ype.
Diagnosis
The new species can be diagnosed from congeners by a combination
of the following morphological characteristics: (1) body length moder-
ate SV L 59.6–67.3 mm; (2) tail short TAL/SVL 141.8%–151.5%; (3) hind
limbs shor t HLL/SVL 60.2%–65.8%; (4) head width moderate HW/HL
69.5%–73.6%; (4) MD 42–46; (5) F4S 14–17; (6) T4S 20–24; (7) conical
or sub -py ramidal post-rictal scale moder ately devel oped, 3–7; (8) tym-
panum con ce al ed ; (9) nu chal cre st fee bly de ve lo pe d wi th no skin folds;
(10) transverse gular fold present, distinct; (11) ventral scales of head
and body distinctively keeled; (12) ventral head scales and ventrolat-
eral body scales homogeneous in size; (13) distinct gular spots absent
in females, but mosaic Light Sulphur Yellow (Color 93) patterns pre-
sent; (14) dorsolateral stripes strongly jagged , White, Light Buff (Color
2), or Light Sulphur Yellow (91); (15) ventral body uniform Pale Buff
(Color 1); and (16) inner lips, oral cavity, and tongue uniform flesh color.
Comparisons
Because all species of Diploderma are sexually dimorphic, and since
we only have females of D. panchi sp. nov., comparisons are carried
out for females only. Diploderma panchi sp. nov. was confused as
D. flaviceps, but it can be differentiated from the latter species by
having feebly developed nuchal crests with no skin folds in females
(versus erected on well-developed skin folds), a relatively shorter tail
TAL ≤ 151.0% SVL in females (versus ≥191.5%), as well as by the
distinct coloration of gular patterns (Light Sulphur Yellow [Color 93]
blotches versus Jet black [Color 30 0] reticulated stripes), and the ab-
sence of dark rhomb-shaped patterns along dorsal midline in females
(versus presence).
The new species was morphologically most similar to D. brevi-
caudum and D. drukdaypo, with all three species having a dwarf ap-
pearance (i.e., disproportionally short tail and short limbs). However,
the new species can be differentiated from D. brevicaudum by hav-
ing more middorsal scales (MD 42–46 versus 34–40) and the pres-
ence of mosaic gular pat terns in females (versus absence); and from
D. drukdaypo by having distinctively keeled ventral scales (versus
smooth or feebly keeled), distinct coloration of dorsolateral stripes
(Light Yellow Ocher [Color 13] to Pale Buf f [Color 1] versus Medium
Chrome Orange [Color 75]), and presence of mosaic gular patterns in
females (versus absence).
For remaining congeners, D. panchi sp. nov. differs from all by
having a much shorter tail in females (TAL TAL < 151.7% SVL in fe-
males versus >160.0%), shorter hind limbs (HLL ≤ 65.8% SVL versus
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>69.0%), and differential gular patterns (mosaic pattern versus no
gular spots or single distinct gular spot). Specifically, D. panchi sp.
nov. differs from D. fasciatum by the absence of transverse hour-
glass-shaped pattern on mid-dorsum (versus presence) and presence
of dorsolateral stripes (versus absence); from D. batangense, D. ia-
dium, and D. yulongense by differential gular coloration and patterns
in females (Light Sulphur Yellow [93], mosaic patterns versus single
spot, Pale Cyan [Color 157] in D. batangense, Medium Greenish Yellow
[Color 88] in D. iadinum, and Chartreuse [Color 89] in D. yulongense);
from D. chapaense, D. micangshanense, D. varcoae, D. yunnanense, and
all species from oceanic islands (D. brevipes, D. luei, D. makii, D. po-
lygonatum, and D. swinhonis) by the presence of distinct transverse
gular fold (versus absence); from D. swild and D. splendidum by hav-
ing homogeneous ventral head scales (versus heterogeneous); from
D. laeviventre by distinct ivel y keeled vent ral sca le s (versus smooth or
feebly keeled); from D. dymondi, D. slowinskii, and D. varcoae by hav-
ing a concealed tympanum (versus exposed); and from D. vela by dif-
ferential coloration of dorsolateral stripes in females (Light Sulphur
Yellow [Color 93] versus Medium Chrome Orange [Color 75]).
For the new species described here, D. panchi sp. nov. differs
from all but D. flavilabre sp. nov. by having a much short t ail in fe-
males (TAL ≤ 151.0% SVL versus ≥174.3%). In addition, D. panchi sp.
nov. differs from D. angustelinea sp. nov. by having more conical or
sub- py r ami da l pos t-ri c tal sc al e s (3–7 ver sus 0–3 ) and stro ngl y ja g ge d
and wide dorsolateral stripes in females (versus feebly jagged and
thin); from D. aorun sp. nov. by di s tin c t gul ar co lor ati o n (L igh t Sul p hu r
Yellow [Color 93], mosaic patterns versus single spot, Pale Cyan
[Color 157] to Light Caribbean Blue [Color 163]). For D. flavilabre sp.
nov. , D. panchi sp. nov. differs by having a distinct gular pattern and
coloration (Light Sulphur Yellow [Color 93], mosaic pat terns versus
Pale Emerald Green [Color 141] to Light Turquoise Green [Color 146]
gular spots) and a distinct lip coloration (Light Flesh Color [Color
250] versus Dark Spectrum Yellow [Color 78]).
Description of holotype
Adult female, body slightly compressed dorsally, SVL 67.3 mm. Tail
slender, short, TAL 151.5% SVL. Limbs short, FLL 40.0% SVL, HLL
61.2% SVL. Head slightly compressed dorsally, HW 71.5% HL, HD
68.8% HW; snout pointy, SEL 37% HL. Rostral rectangular, two scales
away from nasal; nasal oval shape, single scale away from first suprala-
bial; supralabial 9/8, keeled; suborbital scale rows 4/4, distinctively
keeled, third row from bottom enlarged; numerous enlarged scales
between posterior orbit and tympanum. Tympanum concealed under
fine scales; enlarged, conical, or sub-pyramidal scales post-tympanic,
4/5. Sub-pyramidal, post-rictal scales moderately developed, 5/6.
Dorsal head scales heterogeneous, distinctively keeled; Y-shaped
FIGURE 12 Comparisons of lateral head view (1), dorsal head view (2), ventral head view (3), dorsolateral overview (4), and ventral
overview (5) of preser ved specimens among females of Diploderma angustelinea sp. nov. (a), D. aorun sp. nov. (b), D. flavilabre sp. nov. (c),
D. qilin sp. nov. (d), and D. panchi sp. nov. (e). Photographs by KW
(a1) (a2) (a3) (a4) (a5)
(b1) (b2) (b3) (b4) (b5)
(c1) (c2) (c3) (c4) (c5)
(d1) (d2) (d3) (d4) (d5)
(e1) (e2) (e3) (e4) (e5)
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ridge present on dorsal snout, formed by enlarged, modified scales,
from two small scales posterior to rostral extending across forehead
to approximately three scales anterior of interparietal. Interparietal
enlarged, tear shape, distinct parietal eye present. Single, elongated,
subpyramidal scale present on each side post-orbit dorsally. Enlarged,
conical or subpyramidal scales post-temporal, 4/2, with outer most
one twice as large as adjacent one.
Dorsal body scales heterogeneous in size, with enlarged, modi-
fied scales scattered across. Nuchal crest feebly developed with no
skin folds, indistinguishable from dorsal crest; crest scales serrated,
slightly erected, larger than nearby dorsal scales. Two lateral rows
of enlarged scales forming dorsolateral ridges, parallel to vertebral
ridge on each side of body from neck to pelvis. Dorsal limb scales
distinctively keeled, homogeneous on forelimbs, heterogeneous
on hind limbs, with enlarged scales in single oblique row on thigh
and crus, respectively; enlarged conical scales present on posterior
thigh. Tail scales homogeneous, distinctively keeled, craniate in lat-
eral rows.
Ventral head scales homogeneous, distinctively keeled; men-
tal pentagonal in shape, enclosed by first pair of chin shields; chin
shields keeled, 6/6, separated from infralabials by two rows of small
scales. Gular pouch present in life, moderately developed, indistinct
after preservation; transverse gular fold present, distinct. Ventral
body, limb scales homogeneous, distinctively keeled. Finger I short-
est, Finger IV longest, F4S 16/14, T4S 20/20.
Coloration of holotype in life
The dorsal sur face of the head is mostly uniform Cinnamon-Drab
(Color 50), with some unclear Light Buff (Color 2) transverse streaks
between orbits. The background coloration of the lateral surfaces
of the head is Light Buff (Color 2). Burnt Umber (Color 48) to Dark
Neutral Gray (Color 299) radial stripes are present around eyes,
whereas the stripe from posterior–inferior corner of the eye to the
rictus and the one from posterior–superior corner of the eye to the
enlarged conical scale post-supraciliaries are the darkest and long-
est. Rem aining ra di al stri pe s that are infe ri or to the orb it are unclea r
and faint. The background coloration of the lateral head is Light
Buff (Color 2). Light Neutral Gray (Color 297) marble patterns are
present on the loreal region of head, giving a dirty appearance. The
area of lateral head that are in bet ween and posterior to the two
darkest radial stripes and anterior to t ympanum are Light Russet
Vinaceous (Color 246). Scales coving tympanum are Fawn Color
(Color 258).
Background coloration of dorsal and lateral surfaces of limbs,
body, and tail are Cinnamon-Drab (Color 50). A strongly Jagged, zig-
zag- sha pe d , do r s o l a ter a l st r i p e is present on ea ch side of th e body fro m
neck to base of tail, with the most anterior one third of dorsolateral
stripe faint Light Yellow Ocher (Color 13), and the remaining parts of
the stripe Pale Buff (Color 1). The boarding regions along the superior
and inferior edge of dorsolateral stripes are Raw Umber (Color 280),
par tic ularly in the valleys region of the zigzag dorsolateral stripe along
th e su per ior sid e of the dors ola te r al st rip e. The da rk colo rat io n of each
valley along each dorsolateral stripe forms a dark patch, symmetrical
to th e co rresp on di ng one on th e ot her side of th e bo dy. Area alo ng the
FIGURE 13 Habitats of the six new species at their type localities in the Hengduan Mountain Region. (a) Diploderma angustelinea sp.
nov.: near Maidilong Village, Muli County, Liangshan Prefecture, Sichuan Province, China; (b) D. aorun sp. nov.: near Benzilan Township,
Deqin County, Yunnan Province, China; (c) D. flavilabre sp. nov.: near Yebatan Hydropower Station, Baiyu County, Ganzi Prefecture, Sichuan
Province, China; (d) D. panchi sp. nov.: near Yajiang Township, Yajiang Count y, Ganzi Prefecture, Sichuan Province, China; (e) D. panlong sp.
nov.: near Miansha Village, Mianning County, Liangshan Prefecture, Sichuan Province, China; and (f) D. qilin sp. nov.: near Wujin Village,
Shangri-La County, Yunnan Province, China
(a) (b) (c)
(d) (e) (f)
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vertebrate between dorsolateral stripes is dirty Salmon Color (Color
58). Randomly scattered enlarged scales on lateral and ventrolateral
body are Pale Bu ff (Color 1). Co loratio n of la teral bod y gr ad ually fad es
and transition into white inferiorly toward ventral body.
Faint Pale Buff (Color 1) and Salmon Color (Color 58) transverse
streaks are present on the dorsal forelimbs and hind limbs, respec-
tively. A Pale Buff (Color 1), narrow stripe with Raw Umber (Color
280) edges is present from the base of thigh to the knee joint on the
posterior ventrolateral side of hind limb on each side. Four Salmon
Colored (Color 58) triangular patterns with Raw Umber (Color 280)
outlines are present on the dorsal surface from the pelvis to one
fifth of the tail from the vent. The remaining parts of the tail are
patterned with Drab (Color 19) spots or transverse streaks.
The background coloration of the ventral surface of the head is
Pale Buff (Color 1). Faint Jet Black (Color 300) vermiculated stripes
are present on the ventral head, some of which are interconnected
and form a web pattern. Light Sulphur Yellow (Color 93) patches are
present in between the faint Jet Black vermiculate stripes, forming a
mosaic pattern on the gular. Ventral surfaces of the body and limbs
are uniform Pale Buff (Color 1) with no distinct ornamentations, and
the ventral hands and feeds, as well as the ventral surface of the distal
portion of the tail are pale Tawny Olive (Color 17).
Coloration of holotype in preservation
Most ornamentation patterns remain the same after preservation,
except for the Light Sulphur Yellow (Color 93) reticulated patterns of
gular disappear. Additionally, most coloration remains the same except
the Light Yellow Ocher (Color 13) and Pale Buff (Color 1) coloration of
the dorsolateral stripes, which fade to Smoky White (Color 261).
Variatio n
Morphometric and pholidosis variation of the type series is sum-
marized in Table S8. For coloration and ornamentations, all para-
topotypes (KIZ 032717, 032716, and 032729) have reduced black
vermiculated patterns on the ventral surface o the head, and para-
type KIZ 032717 has the most reduced patterns.
Etymology
The species name “panchi” is derived from the name of female drag-
ons (Chinese蟠螭, Pinyin: Pan Chi) in the ancient Chinese literature,
《汉书 ·司马相如传》. We recommend Panchi Mountain Dragon as its
English common name and 蟠螭龙蜥(Pinyin: Pan Chi Long Xi) as its
Chinese common name.
Natural Histor y and conservation
Diploderma panchi sp. nov. in h abi t s the wa r m–d r y vall e y al ong the up p e r
Yalong River, which is predominantly coniferous forest along with tall
bush species (i.e., Lonicera tangutica; Figure 13d). The new species is
terrestrial, basking in the open areas in sunny days and seeking rock
crevices or fallen woodpiles for shelters when disturbed. Individuals
were observed to sleep on bushes at night. Lizard-feeding snakes such
as Elaphe carinata may represent predators of the new species.
Based on our surveys, habitats of the new species overlap greatly
with human inhabitant s, township developments, and hydropower
developments (i.e., Lianghekou Station) in Yajiang County, Sichuan
Province. Although the exact distribution range of the new species
is still unknown, its distribution is unlikely to expand much far ther
away from the type localit y in Yajiang County. We recommend Data
Deficient (DD) as its IUCN assessments, and we call for future eco-
logical studies on the population size and distribution range of the
new species.
3.3.5 | Diploderma panlong sp. nov. Wang, Che, Siler
(Population 8 in Figure 1; Figures 2, 9, 11; Tables 2–4;
Table S8 (5); ZooBank ID urn:lsid:zoobank.org:act:79DA84FF-
CA00- 4C19-9D7D-3F 3071C2AC8F) .
Synonyms
Japalura dymondi Deng, Yu, and C ao (1991: 27), in part. Zhao
et al. (1999: 110–111), in part. Zhao (20 03: 82–83), in par t.
Japalura flaviceps Deng et al. (1991: 27); Zhao et al. (1999: 111–
115); Zhao (2003: 84), in part.
Holotype
KIZ 040138, adult male collected near Siyinuo Village, Mianning
County, Liangshan District, Sichuan Province, China (101.8806° E,
28.3569° N, elevation 1,430 m, WGS 84). Collected by Kai Wang
and Gadeng Nima on April 25, 2018.
Paratopotypes
KIZ 040137, 040139, 04 0140, 040143, adult males; 040141,
040142, young adult males. All share same collecting information
as the holotype.
Diagnosis
The new species can be diagnosed from congeners by a combination
of following morphological characters (1) body size moderate, SVL
60.2–71.7 in males; (2) tail long, TAL 248.2%–268.0% SVL in males;
(3) hind limbs moderate, HLL 77.7%–83.2% SVL; (4) head moder-
ate, HW 63.4%–67.3% HL; (5) MD 39–46; (6) F4S 18–24; (7) T4S
22–27; (8) conical, post-rictal scales strongly developed, 9–13; (9)
tympana mostly exposed; (10) nuchal crest moderately developed,
TNC 5.4%–7.7% HL; (11) transverse gular fold present in life, shal-
low, sometimes indistinct after preservation; (12) ventral head and
body scales homogeneous, distinctively keeled; (13) gular spots ab-
sent in either sexes; (14) dorsolateral stripes smooth-edged, narrow,
Sulphur Yellow (Color 80); (15) distinct radial stripes present around
eyes except suborbital regions; (16) distinc t white lip stripe present
on each side below eye; (17) gradual transition from Tawny (Color
60) or Amber (Color 51) dorsolateral body coloration to Light Flesh
(Color 250) coloration of ventrolateral body surface and eventually
to Light Buf f (Color 2) coloration of ventral body surface; and (18)
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oral cavity Dark Spectrum Yellow (Color 78), tongue Light Chrome
Orange (Color 76).
Comparisons
The new species is most similar to D. swild, with both species hav-
ing exposed tympana (in most individuals of D. panlong sp. nov.),
smooth dorsolateral stripes, and Dark Spectrum Yellow (Color 78)
to Light Chrome Orange (Color 76) oral coloration, and both spe-
cies are found along the Yalong River Valley. However, D. panlong
sp. nov. differs from D. swild by having a relatively longer tail in
males (TAL 248.2%–268.0% SVL versus 225.5%–239.0