A New Species of Diploderma (Reptilia: Squamata: Agamidae) from the upper Salween River in Eastern Tibet, China

Abstract

A new species of Diploderma is described from the upper Salween River Valley in eastern Tibet, China based on morphological and genetic data. The new species is morphologically most similar and phylogenetically closely related to D. laeviventre, but it can be easily diagnosed by having distinct conical scales on the post rictal region of the head, distinctively keeled ventral head and body scales, and different coloration of gular spots and dorsolateral stripes in both sexes. The taxonomic discovery further highlights the underestimated diversity of the genus and the importance of habitat conservation of the neglected hot-dry valley ecosystems in the Hengduan Mountain Region of China.

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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by J. Smid: 21 Jan. 2022; published: 11 Feb. 2022 201
Zootaxa 5099 (2): 201–220
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5099.2.3
http://zoobank.org/urn:lsid:zoobank.org:pub:DDC8A093-1765-4670-B95E-A010DFEEEB52
A New Species of Diploderma (Reptilia: Squamata: Agamidae) from the upper
Salween River in Eastern Tibet, China
KAI WANG1*, YINPENG ZHANG2 & XIANQI LI3
1Sam Noble Oklahoma Museum of Natural History and Department of Biology, University of Oklahoma, Norman, Oklahoma, USA,
73072.
2Department of Zoology, Michigan State University, East Lansing, Michigan, USA, 48823.
�
zhan1198@msu.edu; http://orcid.org/0000-0001-6857-7445
3Key Laboratory of Conserving Wildlife with Small Populations in Yunnan, Southwest Forestry University, Kunming, Yunnan, China,
650224.
�
wolleyil@163.com; https://orcid.org/0000-0002-5396-0983
*Corresponding author.
�
kai.wang-2@ou.edu; https://orcid.org/0000-0002-6736-3346
Abstract
A new species of Diploderma is described from the upper Salween River Valley in eastern Tibet, China based on
morphological and genetic data. The new species is morphologically most similar and phylogenetically closely related
to D. laeviventre, but it can be easily diagnosed by having distinct conical scales on the post rictal region of the head,
distinctively keeled ventral head and body scales, and different coloration of gular spots and dorsolateral stripes in both
sexes. The taxonomic discovery further highlights the underestimated diversity of the genus and the importance of habitat
conservation of the neglected hot-dry valley ecosystems in the Hengduan Mountain Region of China.
Key words: Cryptic diversity, Draconinae, IUCN, Lizard, Vulnerable
Introduction
Mountain Dragons of the genus Diploderma are the most diverse genus of lizards in China, including 33 species
currently recognized from China (Liu et al. 2020; Wang et al. 2019a, 2019b, 2019c, 2020, 2021a, 2021b). In China,
the Hengduan Mountain Region represents the diversity hotspot for Diploderma, and most species are micro-en-
demic and only found in a specific section of a given river valley (Wang et al. 2021a, 2021b). Previous taxonomic
studies of the genus have focused largely on the Mekong (Wang et al. 2015, 2016, 2019b), Yangtze (Manthey et al.
2012; Wang et al. 2017, 2021a, 2021b), and Yalong River Valleys (Wang et al. 2019a, 2019c, 2021a) in the Heng-
duan Mountain Region, and only a few examined the diversity along the Salween River (Wang et al. 2016; Rao et
al. 2017).
Along the Salween River, only two species of Diploderma are currently recognized, namely D. laeviventre
(Wang, Jiang, Siler, Che, 2016) from the far upper reaches in Basu County, Tibet, and D. slowinskii (Rao, Vindum,
Ma, Fu, Wilkinson, 2017) from lower reaches in Gongshan County, Yunnan Province (Fig. 1). As the diversity of the
genus shows similar turnover across river valleys in the Hengduan Mountain Region (Wang et al. 2019b, 2021b),
the large gap along the Salween River between the ranges of D. laeviventre and D. slowinskii may harbor additional
undescribed diversity.
During herpetological surveys in 2020, specimens of the genus Diploderma were collected from the gap region
along the Salween River Valley in Cawarong (=Chawalong) Village, Zayu County, Tibet, China. Combining mor-
phological and mitochondrial genetic data, we here describe them as a new species.

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202 · Zootaxa 5099 (2) © 2022 Magnolia Press
Materials and methods
Sampling
A total of 10 specimens of the putative new species were collected along the Salween River Valley in Cawarong
Village, Zayu County, Tibet, China on 25 June 2019 (Fig. 1). After euthanasia, tissue samples (liver or muscle) were
taken and stored in 95% ethanol, while the specimens were vouchered in 10% buffered formalin and transferred to
75% ethanol after fieldwork. Specimens were deposited at the Southwest Forestry University (SWFU), China.
FIGURE 1. Distribution of the genus Diploderma on mainland East Asia. Star: type locality of Diploderma yangi sp. nov. in
Cawarong (=Chawalong), Zayu County, Tibet, China. Circles: type localities of recognized congeners, which are the followings:
1. D. splendidum, 2. D. micangshanense, 3. D. zhaoermii, 4. D. grahami, 5. D. flaviceps, 6. D. panchi, 7. D. bowoense, 8. D.
angustelinea, 9. D. panlong, 10. D. swild, 11. D. dymondi, 12. D. varcoae, 13. D. flavilabre, 14. D. batangense, 15. D. formos-
gulae, 16. D. aorun, 17. D. qilin, 18. D. yulongense, 19. D. brevicauda, 20. D. drukdaypo, 21. D. vela, 22. D. iadinum, 23. D.
laeviventre, 24. D. slowinskii, 25. D. yunnanense, 26. D. menghaiense, and 27. D. chapaense.
In addition to the newly collected specimens, we also examined specimens of 30 recognized congeners from
the following natural history museum collections: American Museum of Natural History, U.S.A. (AMNH); Natural
History Museum, U.K. (BMNH); California Academy of Sciences, U.S.A. (CAS); Kunming Institute of Zoology,
Chinese Academy of Sciences (KIZ); National Museum of Natural History, U.S.A. (USNM); Museum of Compara-
tive Zoology, U.S.A. (MCZ); Museum of Natural History Vienna, Austria (NMW); and Museum für Naturkunde,
Leibniz Institute for Evolution and Biodiversity Science (ZMB) (Appendix I).
Molecular data and phylogenetic analyses
Total genomic DNA was extracted from liver or muscle tissues of four specimens with a standard three-step phe-
nol-chloroform extraction method. The mitochondrial gene NADH dehydrogenase subunit 2 (ND2) was targeted
and amplified using published primers (Jap_70F, Jap_1559R; Wang et al. 2019a). PCR and sequencing methods
followed recent published protocols (Wang et al. 2021a).
In addition to the newly generated sequences, matching ND2 data for 31 species of Diploderma and represen-

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tatives of Calotes, Pseudocalotes, and Acanthosaura were obtained from GenBank (Table 1). Outgroup taxa were
chosen based on the recent phylogenetic study of the subfamily Draconinae (Wang et al. 2019a). All sequences were
edited and aligned using MUSCLE in Geneious v. 10.0.6. The final alignment contained 1031 bp, with four novel
sequences deposited in GenBank (Table 1).
TABLE 1. GenBank accession numbers for the sequences used. New sequences are indicated in bold.
Vouchered Number Species Locality ND2 GenBank
Accession
Number
SWFU 005410 D. yangi sp. nov. Zayu, Tibet, China OL449603
SWFU 005412 D. yangi sp. nov. Zayu, Tibet, China OL449604
SWFU 005414 D.yangi sp. nov. Zayu, Tibet, China OL449605
SWFU 005411 D. yangi sp. nov. Zayu, Tibet, China OL449606
KIZ 029704 D. angustelinea Muli, Sichuan, China MT577930
KIZ 029705 D. angustelinea Muli, Sichuan, China MT577924
KIZ 029708 D. angustelinea Muli, Sichuan, China MT577931
KIZ 029710 D. angustelinea Muli, Sichuan, China MT577927
KIZ 032488 D. angustelinea Muli, Sichuan, China MT577925
KIZ 032489 D. angustelinea Muli, Sichuan, China MT577926
KIZ 032490 D. angustelinea Muli, Sichuan, China MT577928
KIZ 032491 D. angustelinea Muli, Sichuan, China MT577929
KIZ 032733 D. aorun Benzilan, Yunnan, China MT577938
KIZ 032734 D. aorun Benzilan, Yunnan, China MT577939
KIZ 032735 D. aorun Benzilan, Yunnan, China MT577937
KIZ 032736 D. aorun Benzilan, Yunnan, China MT577936
KIZ 032737 D. aorun Benzilan, Yunnan, China MT577940
KIZ 019278 D. batangense Zhubalong, Markam, Tibet, China MT577932
KIZ 019279 D. batangense Zhubalong, Markam, Tibet, China MT577933
KIZ 019276 D. batangense Batang, Sichuan, China MK001413
KIZ 09404 D. batangense Zhubalong, Tibet, China MK001412
KIZ 019281 D. batangense Batang, Sichuan, China MT577934
KIZ 019314 D. batangense Zhubalong, Markam, Tibet, China MT577935
KIZ 044304 D. brevicauda Lijiang, Yunnan, China MW506023
KIZ 044305 D. brevicauda Lijiang, Yunnan, China MW506021
KIZ 044306 D. brevicauda Lijiang, Yunnan, China MW506022
KIZ 044757 D. bowoense Muli, Sichuan, China MW506020
KIZ 044758 D. bowoense Muli, Sichuan, China MW506019
NMNS 19607 D. brevipes Taiwan, China MK001429
NMNS 19608 D. brevipes Taiwan, China MK001430
KIZ 034923 D. chapaense Lvchun, Honghe, Yunnan, China MG214263
KIZ 040145 D. chapaense Dali, Yunnan, China MK578667
KIZ 046954 D. chapaense Jingdong, Yunnan, China MK578660
KIZ 046970 D. chapaense Jingdong, Yunnan, China MK578659
KIZ 047085 D. chapaense Jingdong, Yunnan, China MK578661
KIZ 034921 D. chapaense Lvchun, Yunnan, China MG214264
ZMMU NAP-01911 D. chapaense Chapa, Vietnam MG214262
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204 · Zootaxa 5099 (2) © 2022 Magnolia Press
TABLE 1. (Continued)
Vouchered Number Species Locality ND2 GenBank
Accession
Number
KIZ 027627 D. drukdaypo Jinduo, Chamdo, Tibet, China MT577950
KIZ 027628 D. drukdaypo Zhuka, Chamdo, Tibet, China MT577592
KIZ 027629 D. drukdaypo Zhuka, Chamdo, Tibet, China MT577593
KIZ 016486 D. drukdaypo Chamdo City, Tibet, China MT577951
KIZ 027630 D. drukdaypo Zhuka, Chamdo, Tibet, China MT577954
KIZ 040147 D. dymondi Panzhihua, Sichuan, China MT577899
KIZ 040148 D. dymondi Panzhihua, Sichuan, China MT577900
KIZ 040149 D. dymondi Panzhihua, Sichuan, China MT577901
KIZ 040639 D. dymondi Dongchuan, Yunnan, China MK001422
KIZ 040640 D. dymondi Dongchuan, Yunnan, China MK001423
KIZ 019575 D. flaviceps Kangding, Sichuan, China MT577896
KIZ 019576 D. flaviceps Kangding, Sichuan, China MT577897
KIZ 019577 D. flaviceps Kangding, Sichuan, China MT577895
KIZ 019578 D. flaviceps Kangding, Sichuan, China MT577894
KIZ 019579 D. flaviceps Kangding, Sichuan, China MT577898
KIZ 01851 D. flaviceps Luding, Sichuan, China MK001416
KIZ 01852 D. flaviceps Luding, Sichuan, China MK001417
KIZ 032692 D. flavilabre Baiyu,Sichuan, China MT577916
KIZ 032694 D. flavilabre Baiyu, Sichuan, China MT577917
KIZ 032695 D. flavilabre Baiyu, Sichuan, China MT577918
KIZ 032696 D. flavilabre Baiyu, Sichuan, China MT577919
KIZ 032697 D. flavilabre Baiyu, Sichuan, China MT577915
KIZ 032698 D. flavilabre Baiyu, Sichuan, China MT577920
KIZ 044420 D. formosgulae Deqin, Yunnan, China MW506024
KIZ 044421 D. formosgulaeDeqin, Yunnan, China MW506025
KIZ 044373 D. formosgulae Deqin, Yunnan, China MW506028
KIZ 044375 D. formosgulae Deqin, Yunnan, China MW506026
KIZ 044376 D. formosgulae Deqin, Yunnan, China MW506027
KIZ 027697 D. iadinum Yunling, Deqin, Yunnan, China MT577956
KIZ 027702 D. iadinum Yunling, Deqin, Yunnan, China MT577957
KIZ 027706 D. iadinum Yunling, Deqin, Yunnan, China MT577955
KIZ 027691 D. laeviventre Basu, Markam, Tibet, China MT577892
KIZ 027692 D. laeviventre Basu, Markam, Tibet, China MT577893
KIZ 014037 D. laeviventre Basu, Markam, Tibet, China MK001407
NMNS 19604 D. luei Taiwan, China MK001433
NMNS 19605 D. luei Taiwan, China MK001434
NMNS 19609 D. makii Taiwan, China MK001431
NMNS 19610 D. makii Taiwan, China MK001432
L0030 D. menghaiense Menghai, Yunnan, China MT598655
L0031 D. menghaiense Menghai, Yunnan, China MT598656
L0032 D. menghaiense Menghai, Yunnan, China MT598657
L0033 D. menghaiense Menghai, Yunnan, China MT598658
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TABLE 1. (Continued)
Vouchered Number Species Locality ND2 GenBank
Accession
Number
KIZ 023231 D. micangshanense Xixia, Henan, China MK578664
KIZ 032801 D. micangshanense Shiyan,Hubei, China MK578665
KIZ 032802 D. micangshanense Shiyan,Hubei, China MK578666
KIZ 06850 D. micangshanense Longnan, Gansu, China MK001424
WK-JK 037 D. micangshanense Wenxian, Gansu, China MK578662
WK-JK 038 D. micangshanense Wenxian, Gansu, China MK578663
KIZ 032715 D. panchi Yajiang, Sichuan, China MT577946
KIZ 032716 D. panchi Yajiang, Sichuan, China MT577944
KIZ 032717 D. panchi Yajiang, Sichuan, China MT577947
KIZ 032729 D. panchi Yajiang, Sichuan, China MT577945
KIZ 040137 D. panlong Miansha, Liangshan, Sichuan, China MT577906
KIZ 040138 D. panlong Miansha, Liangshan, Sichuan, China MT577907
KIZ 040139 D. panlong Miansha, Liangshan, Sichuan, China MT577908
KIZ 040140 D. panlong Miansha, Liangshan, Sichuan, China MT577905
KIZ 040141 D. panlong Miansha, Liangshan, Sichuan, China MT577909
KIZ 040143 D. panlong Miansha, Liangshan, Sichuan, China MT577904
NMNS 19598 D. polygonatum Taiwan, China MK001427
NMNS 19599 D. polygonatum Taiwan, China MK001428
KIZ 028332 D. qilin Balong, Deqin, Yunnan, China MT577941
KIZ 028335 D. qilin Balong, Deqin, Yunnan, China MT577943
KIZ 028333 D. qilin Balong, Deqin, Yunnan, China MT577942
KIZ 027543 D. slowinskii Fugong, Gongshan, Yunnan, China MT577910
KIZ 027544 D. slowinskii Fugong, Gongshan, Yunnan, China MT577911
KIZ 027572 D. slowinskii Qiunatong, Gongshan, Yunnan, China MT577912
KIZ 027573 D. slowinskii Qiunatong, Gongshan, Yunnan, China MT577913
CAS 214906 D. slowinskii Gongshan, Yunnan, China MK001405
CAS 214954 D. slowinskii Gongshan, Yunnan, China MK001406
KIZ 015973 D. splendidum Yichang, Hubei, China MK001418
CAS 194476 D. splendidum Yaan, Sichuan, China AF128501
KIZ 034893 D. swild Panzhihua, Sichuan, China MN266297
KIZ 034894 D. swild Panzhihua, Sichuan, China MN266300
KIZ 034895 D. swild Panzhihua, Sichuan, China MN266298
KIZ 034914 D. swild Panzhihua, Sichuan, China MN266299
KIZ 034915 D. swild Panzhihua, Sichuan, China MN266301
NMNS 19592 D. swinhonis Taiwan, China MK001419
NMNS 19593 D. swinhonis Taiwan, China MK001420
KIZ 029711 D. varcoae Dali, Yunnan, China MT577902
WK-JK 011 D. varcoae Yuxi, Yunnan, China MT577903
KIZ 026132 D. varcoae Mengzi, Honghe, Yunnan, China MK001421
KIZ 027672 D. vela Tongsha, Markam, Tibet, China MT577949
KIZ 027673 D. vela Tongsha, Markam, Tibet, China MT577948
KIZ 034925 D. vela Quzika, Markam, Tibet, China MK001415
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206 · Zootaxa 5099 (2) © 2022 Magnolia Press
TABLE 1. (Continued)
Vouchered Number Species Locality ND2 GenBank
Accession
Number
KIZ 019299 D. vela Quzika, Markam, Tibet, China MK001414
KIZ 028291 D. yulongense Shangrila, Yunnan, China MT577921
KIZ 028292 D. yulongense Shangrila, Yunnan, China MT577922
KIZ 028300 D. yulongense Shangrila, Yunnan, China MT577923
KIZ 09399 D. yulongense Xianggelila, Yunnan, China MK001410
KIZ 43196 D. yulongense Xianggelila, Yunnan, China MK001411
KIZ 040193 D. yunnanense Yingjiang, Yunnan, China MT577914
KIZ 040193 D. yunnanense Yingjiang, Yunnan, China MK578658
CAS 242271 D. yunnanense Baoshan, Yunnan, China MK001408
CAS 242183 D. yunnanense Baoshan, Yunnan, China MK001409
KIZ 019564 D. zhaoermii Wenchuan, Sichuan, China MK001425
KIZ 019565 D. zhaoermii Wenchuan, Sichuan, China MK001426
MVZ 216622 D. zhaoermii Wenchuan, Sichuan, China AF128500
–Acanthosaura lepidogaster – AF128499
CAS 223063 Calotes emma – DQ289460
CIB 091468 C. versicolor Hainan, China KC875820
MVZ 224106 Pseudocalotes brevipes Vinh Phuc, Vietnam AF128499.1
TNHC 58040 P. flavigula Perak, Malaysia AF128503.1
CAS 241966 P. kingdonwardi Dulongjiang, Yunnan, China MK001436
CAS 242628 P. kingdonwardi Dulongjiang, Yunnan, China MK001437
KIZ 015975 P. kakhiensis Gongshan, Yunnan, China MK001435
Bayesian inference and Maximum Likelihood analyses were conducted on the final alignment. Bayesian analy-
sis was done using the program MrBayes v. 3.2.6 (Ronquist et al. 2012) on CIPRES. The ND2 coding region was
partitioned by codon position. Using JMODELTEST2 v. 2.1.10 (Guindon & Gascuel 2003; Darriba et al. 2012), the
best evolutionary model of nucleotide substitution was selected for each partition, which inferred the model GTR +
Γ for all partitions. Two independent Markov Chain Monte Carlo analyses were run, each with four Metropolis-cou-
pled chains, a melting temperature of 0.02, and an exponential distribution with a rate parameter of 25 as the prior
on branch lengths (Marshall 2010). Each run was conducted with 90 million generations and sampled every 2,000
generations, discarding the first 20% of trees as burnin. Finally, we confirmed convergence of runs with TRACER
v. 1.6.0 (visual convergence and Effective Sampling Size>200; Rambaut et al. 2013).
Maximum Likelihood analysis was conducted using RAxML-VI-HPC v. 8.2.10 (Stamatakis, 2014). The most
complex model (GTR + Γ) was applied to all partitions, with 1,000 replicate Maximum Likelihood inference runs.
We initiated each inference with a random starting tree and assessed nodal support with 1,000 bootstrap pseudorep-
licates (Stamatakis et al. 2008). Finally, uncorrected genetic pairwise distances (p-distances) were obtained for the
coding region of ND2 (1031 bp, including <31 bp of missing data on either of the two ends) using PAUP v. 4.0b10
(Swofford 2003).
Morphological data
Specimens were measured using a digital caliper to the nearest 0.1 mm, except for tail length, which was measured
to the nearest 1 mm using a ruler. The following morphometric characters were measured following Wang et al.
(2021a) (abbreviations given in parentheses): snout–vent length (SVL): from the snout tip to anterior edge of the
cloaca; tail length (TAL): from the anterior edge of the cloaca to the tip of tail; head length (HL): from the tip of
snout to the jaw joint; head width (HW): measured between the widest points of the head; head depth (HD): mea-
sured as the perpendicular distance at the temporal region of head; snout–eye length (SEL): measured between the

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tip of snout and anterior edge of orbital bone; fore-limb length (FLL): between the point of insertion at axillary to
the tip of Finger IV, excluding the claw, measured as the limb straighten; hind limb length (HLL): between the point
of insertion at groin to the tip of Toe IV, excluding the claw, measured as the limb straighten; Toe IV length (T4L):
measured between the tip of Toe IV to the base between Toe III and IV, excluding the claw; and trunk length (TRL):
measured between the limb insertion points between axillary and groin. In addition, the following pholidosis char-
acters were recorded: supralabial scale count (SL): number of enlarged, modified labial scales from rostral to the
corner of mouth; infralabial scale count (IL): number of enlarged, modified labial scales from mental to the corner of
mouth; middorsal crest scale count (MD): number of modified crest scales longitudinally from the first nuchal crest
to the scale above cloaca; gular scale count (GU): number of gular scales counted in a straight line along medial axis
between and excluding mental and transverse gular fold; ventral scale count (VN): number of ventral body scales
counted in a straight line along the medial axis between the transverse gular fold and the anterior edge of cloaca;
Finger IV subdigital lamellae count (F4S): number of subdigital lamellae scale from the base between Finger III
and IV to the tip of Finger IV, excluding the claw; Toe IV subdigital lamellae count (T4S): number of subdigital
lamellae scales from the base between Toe III and IV to the tip of Toe IV, excluding the claw; nasal–supralabials
scale rows (NSL): number of horizontal rows of small scales between the first supralabial and the nasal; suborbital
scale rows (SOR): number of longitudinal rows of scales between supralabials and inferior-most edge of orbit circle,
excluding fine ciliary scales in the orbit; canthus rostralis scale count (CR): number of elongated, overlapping scales
from posterosuperior nasal to and including last supraciliary scale along the canthal ridge; post-occipital scale count
(POS): number of enlarged, distinctively keeled, raised conical or sub-pyramidal scales on the occipital region of
head; post-tympanic scale count (PTY): number of enlarged, distinctively keeled, raised conical or sub-pyramidal
scales posterior to the tympana and superior to the rictus; and post-rictal scale count (PRS): number of enlarged,
distinctively keeled, raised conical or sub-pyramidal scales posterior to the rictus and inferior to the tympana. Mea-
surements were taken on the left side of the specimen only (unless the left side was damaged, then the right side was
used), and values for paired pholidosis characters (SL, IL, NSL, CR, SOR, PTY, and PRS) were recorded on both
sides of the body, with counts provided in left/right order. In addition, the following morphological characters were
examined: transverse gular fold across the neck (presence or. absence), gular pouch (absence, feeble, or strongly
developed), and tympana state (exposed or concealed under fine scales).
The following coloration and ornamentation patterns were examined following Wang et al. (2021b): coloration
of the oral cavity (CO), defined as the background coloration of the anterior roof and sides of the mouth, excluding
the posterior palate and deep throat; coloration of the tonsil (CT), defined as the coloration of the posterior palate
and tonsil; coloration of the tongue (CTG); coloration of dorsolateral stripes (CDS); and the shape of dorsolateral
stripes (SDS), either smooth edged or jagged. Color names and codes for coloration followed Köhler (2012).
Results
Molecular results
The genus Diploderma is recovered as monophyletic (Bayesian posterior probability 1.00/maximum likelihood
bootstrap support 95, herein given in this order), and relationships of recognized congeners are identical to previous
studies (Wang et al. 2021a, 2021b; Fig. 2). Individuals from Cawarong form a strongly supported monophyletic
group (1.00/100), which is sister to D. laeviventre (1.00/100); and together these two form a strongly supported
monophyletic group with D. slowinskii (1.00/99).
The putative new species is genetically divergent from its sister species D. laeviventre (uncorrected genetic dis-
tance 6.0–6.5%)and D. slowinskii (14.3–15.0%), which are both found lower the same river. Furthermore, it is also
highly divergent from D. flaviceps (19.9–20.9%), with which the Cawarong population was confused (Table 2).

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208 · Zootaxa 5099 (2) © 2022 Magnolia Press
FIGURE 2. Phylogeny of the genus Diploderma inferred from mitochondrial gene ND2 (1031 bp) using both maximum
likelihood and Bayesian analyses. The maximum likelihood bootstrap support values are mapped onto the Bayesian topology,
where “–” indicate differential topology at the node between the two analyses, with discrepancy on the relationships among D.
brevipes, D. luei, and D. makii. Terminal nodes that unify each recognized species are all strongly supported and hence omitted
(1.00/100).

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TABLE 2. Uncorrected genetic distances among samples of Diploderma yangi sp. nov. and closely related congeners.
Species Voucher Number 1 2 3 4 5 6
1D. flaviceps KIZ 01581 –
2D. flaviceps KIZ 01852 1.4% –
3D. slowinskii KIZ 027543 20.2% 19.8% –
4D. slowinskii KIZ 027544 20.1% 19.9% 0.0% –
5D. slowinskii KIZ 027572 20.4% 20.3% 0.7% 0.7% –
6D. slowinskii KIZ 027573 20.4% 20.3% 0.7% 0.7% 0.0% –
7D. laeviventre KIZ 014037 21.2% 20.6% 16.2% 16.1% 15.8% 15.8%
8D. laeviventre KIZ 027691 21.2% 20.6% 16.2% 16.1% 15.8% 15.8%
9D. laeviventre KIZ 027692 20.2% 20.1% 16.3% 16.3% 15.9% 15.9%
10 D. yangi sp. nov. SWFU 005410 20.8% 20.5% 15.0% 14.8% 14.5% 14.5%
11 D. yangi sp. nov. SWFU 005412 20.8% 20.5% 15.0% 14.8% 14.5% 14.5%
12 D. yangi sp. nov. SWFU 005414 20.8% 20.5% 15.0% 14.8% 14.5% 14.5%
13 D. yangi sp. nov. SWFU 005411 20.9% 20.5% 15.0% 14.7% 14.3% 14.3%
TABLE 2. (Continued)
Species Voucher Number 7 8 9 10 11 12 13
1D. flaviceps KIZ 01581
2D. flaviceps KIZ 01852
3D. slowinskii KIZ 027543
4D. slowinskii KIZ 027544
5D. slowinskii KIZ 027572
6D. slowinskii KIZ 027573
7D. laeviventre KIZ 014037 –
8D. laeviventre KIZ 027691 0.0% –
9D. laeviventre KIZ 027692 0.0% 0.0% –
10 D. yangi sp. nov. SWFU 005410 6.4% 6.4% 6.0% –
11 D. yangi sp. nov. SWFU 005412 6.4% 6.4% 6.0% 0.0% –
12 D. yangi sp. nov. SWFU 005414 6.4% 6.4% 6.0% 0.0% 0.0% –
13 D. yangi sp. nov. SWFU 005411 6.5% 6.5% 6.0% 0.0% 0.0% 0.0% –
Morphological results
The Cawarong population is morphologically most similar to D. laeviventre. Both species have smooth dorsolateral
stripes, feeble nuchal crests in males, and speckled gular regions and colorful gular spots in both sexes. However,
the Cawarong population differs from D. laeviventre by having different pholidosis and coloration characters, and it
can be easily distinguished from all remaining recognized congeners by a combination of morphological characters
(see comparison section below for details; Table 3, 4).
Therefore, considering its monophyletic nature, high genetic divergence, and conspicuous morphological diag-
nosis, we describe the Cawarong population as a new species.
Taxonomic account
Diploderma yangi sp. nov.
(Table 3, 4; Fig. 3–6)
ZooBank urn:lsid:zoobank.org:act:83878ED4-046E-4D75-B88E-E1684226BE5A
Chresonyms. Japalura flaviceps Zhao & Yang 1997: 165–167; Zhao et al. 1999: 111–115; Japalura laeviventris Rao et al.
2017.

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Holotype. SWFU 005414, adult male, from Cawarong Village, Zayu County, Tibet Autonomous Region, China
(28.479ºN, 98.4656ºE, elevation 1887 m). Collected by Liu Xiaolong, Li Xianqi, and Tie Minhua on 6 September
2020.
Paratypes. SWFU 005415, 006812, adult males; SWFU 005417, 005418, subadult males; SWFU 005410–13,
005419, adult females. All share the same locality and collector information as the holotype.
Etymology. The new species name is derived from the last name of Chinese herpetologist, Dr. Datong Yang,
from Kunming Institute of Zoology, Chinese Academy of Sciences. We name the new species after Dr. Yang in hon-
oring his pioneer works and lifetime contributions to the herpetological research in the Hengduan Mountain Region
in China. We suggested Zayu Mountain Dragon as its English common name, and 察隅龙蜥 (Pinyin: Chá Yú Lóng
Xî) as its Chinese common name.
Diagnosis. Diploderma yangi sp. nov. can be diagnosed from congeners by a combination of the following mor-
phological characters: (1) adult body size moderate, SVL 57.8–64.5mm in adults; (2) tail long, TAL 202.8–216.6%
SVL in males, 196.8–208.8% in females; (3) head width moderately wide, HW 69.0%–73.9% HL; (4) hind limbs
long, HLL 69.6–86.3% SVL; (5) tympanum concealed; (6) crest scales small and numerous, MD 45–59; (6) feeble
skin fold under nuchal crest in males, no skin fold under dorsal crest; (7) conical scales present on post temporal,
post tympanic, and post rictal regions, each bearing 3–6 keels, POS 2–6, PTY 2–7, PRS 2–6; (8) F4S 14–18, T4S
20–24; (9) NSL mostly 1, rarely 0; (10) transverse gular fold present, distinct; (11) ventral head and body scales
strongly keeled; (12) distinct radial dark stripes present around eyes; (13) dorsal background coloration Pale Pinkish
Buff (Color 3), speckled with Warm Sepia (Color 40) to Dusky Brown (Color 285) spots laterally inferior to dorso-
lateral stripes; (14) dorsolateral stripes smooth edged and Pale Emerald Green (Color 141) to Light Pistachio (Color
101) in males, slightly wavy and Pale Buff (Color 1) to Pinkish Buff (Color 3) in females; (13) gular spot present in
both sexes, Pale Emerald Green (Color 141) to Light Pistachio (Color 101), paler in females; and (14) oral cavity
and tongue pale Light Flesh Color (Color 250).
FIGURE 3. Diploderma yangi sp. nov. in life. A1: holotype male SWFU 005414; A2: paratype female SWFU 005412; B1:
unvouchered topotypic male; B2: unvouchered topotypic female. Photos by Xiaolong Liu and Jiansheng Peng.

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FIGURE 4. Voucher specimen of the male holotype (A: SWFU 005414) and paratype (B: female SWFU 005412) of Diplo-
derma yangi sp. nov.. 1: dorsal view; 2: ventral view. Photos by Yinpeng Zhang.
FIGURE 5. Head closeup views of the male holotype (A: SWFU 005414) and female paratype (B: SWFU 005412) of Diplo-
derma yangi sp. nov. 1: dorsal head view; 2: lateral head view of the right side; and 3: ventral head view. Photos by Yinpeng
Zhang.

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FIGURE 6. Habitat at the type locality of Diploderma yangi sp. nov. near Cawarong Village, Zayu County, Tibet, China. Photo
by Xiaolong Liu.
Comparisons. Diploderma yangi sp. nov. is phylogenetically sister to and morphologically most similar to D.
laeviventre, but D. yangi sp. nov. can be differentiated from the latter by the presence of well-developed conical
scales in the post-tympanic and post-rictal regions of the head (vs. absence), strongly keeled ventral body scales (vs.
smooth), a different coloration of dorsolateral stripes (Pale Emerald Green [Color 141] to Light Pistachio [Color
101] in males, Pale Buff [Color 1] to Pinkish Buff [Color 3] in females vs. Cream Color [Color12] in males, Sulphur
Yellow [Color 80] in females), and a distinct coloration of gular spots (Pale Emerald Green [Color 141] to Light
Pistachio [Color 101] in both sexes vs. Light Chrome Orange [Color76] in both sexes) (Table 3).
Diploderma yangi sp. nov. was confused with D. flaviceps, but the new species differs from the latter by hav-
ing a tendency toward fewer SL (7–9 vs. 9–11), smooth edged, Pale Emerald Green (Color 141) to Light Pistachio
(Color 101) dorsolateral stripes in males in life (vs. strongly jagged, Pale Horn Color [Color 11]), presence of dis-
tinct radial stripes around the eyes (vs. absence), presence of gular spots in both sexes (vs. absence), absence of skin
folds under nuchal crest in females in life (vs. presence), and absence of dark reticulated patterns on gular region
(vs. presence) (Table 3).
For congener that is from the same river valley downstream, D. yangi sp. nov. differs from D. slowinskii by
having concealed tympana under fine scales (vs. exposed), a much smaller body size (maximum body size 64.1
mm vs. 99.5 mm), less developed and less keeled conical scales on post temporal regions of the head (vs. strongly
developed with multiple keels), the presence of gular spots in both sexes (vs. absence in both sexes), and a terrestrial
lifestyle (vs. arboreal) (Table 3).

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TABLE 3. Morphological comparison among Diploderma yangi sp. nov., its closely related parapatric species, and the
species of historical confusion. “–“ indicates not applicable.
Characters Species
D. yangi sp. nov. D. laeviventre D. slowinskii D. flaviceps
Distribution with respect
to D. yangi sp. nov.
– Parapatric Parapatric Allopatric
Conical Scale Post
Tympanum
Present Absent Present Present
Conical Scale Post
Rictus
Present Absent Present Present
Tympanum State Concealed Concealed Exposed Concealed
Nuchal Crest On feeble skin fold in
males only
On feeble skin fold
in males only
No skin fold in either
sexes
On strong skin fold
in both sexes
Ventral Scale State Distinctively keeled Feebly keeled or
smooth
Distinctively keeled Distinctively keeled
Gular Spot Present Present Absent Absent
Gular Spot Color Pale Emerald Green
(Color 141) to Light
Pistachio (Color 101)
Light Chrome
Orange (Color76)
– –
Shape of Dorsolateral
Stripe in Males
Smooth edged Smooth edged Smooth edged Strongly jagged
Color of Dorsolateral
Stripe in Males
Pale Emerald Green
(Color 141) to Light
Pistachio (Color 101)
Cream Color
(Color 12)
Pale Emerald Green
(Color 141) to Light
Pistachio (Color 101)
Cream Color
(Color 12)
Ecotype Terrestrial Terrestrial Arboreal Terrestrial
From the remaining allopatric species in the Hengduan Mountain Region (isolated by snow covered mountains
of over 4000 m; Fig. 1), D. yangi sp. nov. differs from D. drukdaypo, D. dymondi, D. fasciatum, D. micangsha-
nense, D. panlong, D. splendidum, D. swild, D. varcoae, and D. vela by the presence of colorful gular spots in both
sexes (vs. absence); from all but D. aorun, D. batangense, D. flavilabre, D. grahami, D. hamptoni, D. yulongense
and D. zhaoermii (i.e. D. angustelinea, D. bowoense, D. brevicauda, D. chapaense, D. formosgulae, D. iadinum, D.
menghaiense, D. panchi, D. qilin, D. yunnanense) by having a distinct coloration of gular spots (Pale Emerald Green
[Color 141] to Light Pistachio [Color 101] in both sexes vs. blue, yellow, orange, or pinkish red); from D. batan-
gense, D. flavilabre, D. yulongense, and D. zhaoermii by having smooth dorsolateral stripes in males (vs. strongly
jagged); and from D. grahami by having much longer tail (TAL 202.8–216.6% in males, 196.8–208.8% in females
vs. 164.3%) and distinct transverse gular fold (vs. feeble). Additionally, the new species differ from D. brevicauda,
D. drukdaypo, D. flavilabre, D. panchi by having a much longer tail (TAL 202.8–216.6% in males, 196.8–208.8%
in females vs. <190% in males, <170% in females); from D. aorun, D. batangense, D. bowoense, D. formosgulae,
D. qilin, D. yulongense, and D. zhaoermii by having smooth dorsolateral stripes in males (vs. strongly jagged); from
D. iadinum, D. micangshanense, D. varcoae, and D. zhaoermii by having more middorsal crest scale (45–59 vs.
≤43); from D. angustelinea by the presence of distinct black radial stripes around eyes (vs. absence) and different
shape of dorsolateral stripes in males (smooth edged and wide vs. slightly jagged and narrow); from D. chapaense,
D. fasciatum, D. hamptoni, D. menghaiense, D. micangshanense, D. varcoae, D. yunnanense, and all five species
from East Asian Islands (D. brevipes, D. luei, D. makii, D. polygonatum, and D. swinhonis) by the presence of a
transverse gular fold (vs. absence); and from D. dymondi, D. varcoae, D. swild, and D. panlong by having concealed
tympana (vs. exposed).
Description of holotype. Adult male, body not compressed, SVL 55.9 mm; tail long, slender, TAL 121 mm,
216.6% SVL; forelimb 26.2 mm on left, 46.8% SVL; hindlimb 43.6 mm on left, 78.0% SVL. Head relatively robust,
HW 73.9% HL, HD 73.1% HW; SEL 38.1% HL. Rostral rectangular, bordering seven small postrostral scales; dor-
sal head scales heterogeneous, all strongly keeled; single distinct Y-shaped ridge on dorsal head, stem of Y-shaped
ridge consisting of three enlarged, protruding scales along lateral midline of snout from midpoint between nasal

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to midpoint between anterior to edges of first supraciliaries of each side; single enlarged, protruding scale post su-
praciliary on each side, sub-pyramidal in shape, each bearing 3 or 4 keels; POS 6/4, conical in shape, each bearing
2 or 3 keels, one of each side strongly protruding, larger than remaining ones. Nasal rounded, separated from first
supralabial by single row of scales; loreals small, keeled; suborbital scale rows 3/3, keeled; canthus rostralis elon-
gated except last two posteriorly, greatly overlapping with each other, 10/10; enlarged, keeled scales forming single
lateral ridge from posteroinferior eye to posterosuperior tympanum on each side, 8/7; tympana concealed under fine
scales; SL 8/8, feebly keeled. Mental triangular; IL 7/9, first pair not in contact; enlarged chin shields 4/6, weakly
keeled, first one contacting IL on each side, remaining ones separated from IL by single row of small scales; ven-
tral head scales homogeneous in size, strongly keeled; gular scales 29, homogeneous, distinctively keeled, slightly
smaller or equal to ventrals; distinct transverse gular fold present; gular pouch well developed in life, reduced after
preservation.
Distinct shoulder fold present, short; dorsal body scales heterogeneous in size and shape, all keeled, each bear-
ing single lateral keel; axillary scales fine, much smaller than remining dorsals; enlarged dorsal scales forming two
lateral rows from neck to pelvis on each side of dorsal crest: first row two scales away from dorsal crest, second row
along inferior edge of dorsolateral stripe; remaining enlarged dorsal scales randomly scattered. Nuchal crest scale
triangular, larger and differentiated from dorsal crests in shape, TNC 4.8% HL; dorsal crest scale in fan-shape, tip
pointing anteriorly, each bearing single lateral keel, slightly raised posteriorly; feeble skin fold under nuchal crest;
dorsal crest scales low without skin folds; MD 49. Dorsal limb scales strongly keeled, mostly homogeneous, except
a few enlarged, conical scales on postaxial thighs; F4S 16/15, T4S 22/21. Ventral body scales 59, homogeneous, all
distinctively keeled, mostly carinate in regular lateral rows. Ventral limb scales homogeneous, slightly larger than
ventrals, all distinctively keeled, carinate in regular rows along proximal-distal axis. Tail scales all distinctively
keeled, heterogeneous at anterior most 1/5 of length dorsally, with enlarged scales continuing from dorsal crest
along medial axis and along dorsolateral line on each side; 13 enlarged scales aliened roughly along medial axis on
ventral tail five scales posterior to posterior cloaca lip; remaining tail scales homogeneous, strongly keeled, cari-
nated.
Coloration. In life, the dorsal and lateral surfaces of head are Pale Pinkish Buff (Color 3) to Buff (Color 5),
darker on the dorsal side. Clay Color (Color 18) to Sepia (Color 286) radial stripes are present around the eyes, with
the posteroinferior stripe widest and darkest, and the remaining inferior stripes much narrower and fainter. Three
Sepia (Color 286) transverse streaks are present on dorsal surface of the head, with the first one anterior to eyes, the
second one between eyes, and the last one posterior to the eyes. Clay Color (Color 18) to Sepia (Color 286) speckles
and short streaks are present on the remaining areas of dorsal and lateral surfaces of head.
The background coloration of dorsal and lateral surfaces of the body is Beige (Color 254) to Pale Cinnamon
(Color 55). Single Jet Black (Color 300) triangular patch is present along the posterior side of shoulder fold. A dis-
tinct, smooth-edged, Light Pistachio (Color 101) dorsolateral stripe is present from the neck to the pelvis close to
the dorsal crest on each side. Six rectangular or triangular, Raw Umber (Color 23) to Jet Black (Color 300) patches
are evenly scattered from neck to the pelvis along the vertebral line between the dorsolateral stripes. The region
adjacent to the inferior edge of dorsolateral stripes on each side is Raw Umber (Color 23) to Jet Black (Color 300),
which then gradually transitions to the lighter background coloration of lateral body. Raw Umber (Color 23) to Jet
Black (Color 300) speckles are scattered on the lateral surface of body. Dorsal surfaces of limbs and tail are Beige
(Color 254) to Pale Cinnamon (Color 55). Raw Umber (Color 23) to Jet Black (Color 300) transverse bands are
evenly scattered from the proximal to the distal end on limbs, and from anterior to posterior end on the tail.
The ventral surface of the head is uniform white. A triangular, Light Pistachio (Color 101) gular spot is present
on the posterior gular region. Ventral surfaces of the body, limbs, and tail are uniform white or Pale Buff (Color 1)
with no ornamentation patterns. The oral cavity and tongue pale Light Flesh Color (Color 250).
In preservation, the ornamentation patterns remain the same as in life, but the coloration fades away. Specifi-
cally, the background coloration of dorsal and lateral surfaces of head and body become Pale Neutral Gray (Color
296), all the darker ornamentation patterns (i.e. radial stripes around the eyes, transverse bands on dorsal surface of
head and limbs, and rectangular patches along the dorsal midline of body) all become Medium Neutral Gray (298)
to Dark Neutral Gray (Color 299), and the dorsolateral stripes and the gular spot become Light Venetian Blue (Color
165).
Variations. Detailed morphological variation are summarized in Table 4. Sexual dichromatism are evident in
the new species, in which the males have larger gular spots, and smooth, Pale Emerald Green (Color 141) to Light

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TABLE 4. Morphometric measurements and pholidosis counts of the type series of Diploderma yangi sp. nov. All morphometric measurements are in the unit of mm, and all
ratios are in the unit of %. “–“ indicates missing data due to an incomplete tail or toe. All paired pholidosis characters are given as left/right. For details on the abbreviations see
Materials and Methods.
Characters Voucher Numbers
SWFU
005414
SWFU
005415
SWFU
006812
SWFU
005417
SWFU
005418
SWFU
005410
SWFU
005411
SWFU
005412
SWFU
005413
SWFU
005419
Type Status Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
Sex M M M M JM F F F F F
SVL 55.9 60.0 54.1 47.9 45.4 60.0 57.8 59.6 63.2 64.1
TAL 121.1 – 109.9 102.6 92.2 125.2 – 119.7 124.3 129.4
HW 11.8 12.7 10.6 9.8 9.6 13.1 12.3 12.1 13.3 12.8
HL 15.9 17.8 14.7 14.2 13.2 18.0 17.1 16.6 18.3 18.2
HD 8.6 9.1 7.8 7.4 7.3 9.7 8.8 8.9 9.3 9.1
SEL 6.1 7.4 6.7 6.2 5.7 7.4 7.5 6.9 7.7 7.9
TNC 0.8 1.0 0.6 0.5 0.4 0.6 0.6 0.6 0.7 0.7
FLL 26.2 29.1 26.4 22.5 23.2 24.8 28.8 29.0 27.1 30.9
HLL 43.6 51.8 43.3 40.5 34.5 47.0 48.6 45.6 44.0 47.9
T4L 11.7 12.4 11.1 9.9 9.2 11.5 11.7 12.0 10.4 12.0
TRL 27.8 27.7 24.7 23.4 22.5 27.1 28.6 30.1 32.0 29.9
TAL/SVL (%) 216.6 – 203.2 214.2 202.8 208.8 – 200.7 196.8 202.0
HW/HL (%) 73.9 71.4 72.2 69.0 72.4 72.9 72.2 72.7 72.9 70.7
HD/HL (%) 54.0 51.3 52.7 52.0 54.9 53.9 51.5 53.5 50.8 50.2
HD/HW (%) 73.1 71.9 73.0 75.3 75.8 73.9 71.3 73.6 69.7 71.0
SEL/HL (%) 38.1 41.4 45.3 43.3 43.0 41.1 44.0 41.6 42.2 43.5
FLL/SVL (%) 46.8 48.5 48.8 47.0 51.0 41.3 49.9 48.7 42.9 48.3
HLL/SVL (%) 78.0 86.3 80.1 84.5 75.9 78.3 84.1 76.5 69.6 74.8
TRL/SVL (%) 49.6 46.1 45.7 48.8 49.5 45.2 49.4 50.5 50.7 46.7
TNC/HL (%) 4.8 5.8 4.3 3.5 3.2 3.1 3.4 3.3 3.7 4.0
SL 8/8 9/9 8/8 7/7 8/9 9/9 7/8 8/8 8/7 8/7
IL 7/9 9/10 9/9 8/8 9/9 9/9 9/9 9/9 8/8 9/9
CR 10/10 9/9 9/9 9/9 12/10 9/10 9/8 10/9 10/9 9/8
......continuied on the next page

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TABLE 4. (Continued)
Characters Voucher Numbers
SWFU
005414
SWFU
005415
SWFU
006812
SWFU
005417
SWFU
005418
SWFU
005410
SWFU
005411
SWFU
005412
SWFU
005413
SWFU
005419
Type Status Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
Sex M M M M JM F F F F F
NSL 1/1 1/1 1/1 0/0 1/1 1/1 1/1 1/1 1/1 1/1
SOR 3/3 3/3 3/3 3/3 3/3 3/3 3/3 3/3 3/3 3/3
MD 49 58 49 45 59 47 46 54 45 52
GU 29 27 28 32 31 25 29 26 28 30
VT 59 57 61 58 61 54 56 64 59 59
F4S 16/15 18/16 15/– 18/16 17/17 18/18 16/17 18/17 15/14 18/17
T4S 22/21 23/23 20/21 21/23 22/22 24/24 21/22 22/– 22/23 22/–
POS 6/4 2/3 6/3 3/3 3/3 4/4 5/4 4/5 2/4 4/3
PTY 3/5 3/7 4/2 5/6 4/2 3/4 4/4 5/3 5/6 4/3
PRS 6/2 3/5 6/5 4/3 4/4 6/4 5/4 3/5 2/4 3/4

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Pistachio (Color 101) dorsolateral stripes (vs. shorter tail, smaller gular spots, and irregular and wavy, Pale Buff
[Color 1] to Pinkish Buff [Color 3] dorsolateral stripes in females). The male holotype has a much longer tail than
the females (TAL 216.6% SVL vs. 196.8–208.8% in females, n = 4), but given the limited sample size of adult
males, we cannot conclude on the sexual dimorphism with confidence.
Natural history and conservation. Diploderma yangi sp. nov. is known from the upper Salween River Val-
ley in Zayu County, Tibet Autonomous Region, China. While the upper distribution limit of the new species with
respect to D. laeviventre is unknown, the lower distribution range of the new species is at the Tibet-Yunnan border
region. Interestingly, the distribution border between D. yangi sp. nov. and D. slowinskii matches the climatic turn-
over between hot-dry valley climate and subtropical moist climate along the upper Salween River, and the climatic
factor might have initiated the speciation between the two species and kept them from hybridization. Future ecologi-
cal and evolutionary studies are needed to test this hypothesis.
The species is terrestrial, inhabiting the shrublands of the hot-dry Salween Valley (Fig. 5). Population density
was high, and individuals were observed resting under thick spiky bushes such as Sophora davidii, which were
about 1.5m tall. Sympatric reptiles include Elaphe taeniura and Lycodon gongshan of the family Colubridae and
Gloydius lipipengi of the family Viperidae, and the last two lizard-eating species may be the main predator of D.
yangi sp. nov.. None of the collected females were gravid, suggesting that the survey time in September already
passed the breeding season of the species. At the time of collection, the habitat at the type locality was still in good
condition with no major constructions, but given the overlap between the habitat of the new species and human
habitations along the upper Salween River, expected future expansion of township and infrastructures may pose
threats to the species.
Discussion
Recent taxonomic studies of the genus Diploderma have doubled its diversity from 17 species in 2002 to 35 species
to date (Manthey et al. 2012; Wang et al. 2015, 2016, 2019b, 2019c, 2021a, 2021b; Rao et al. 2017; Liu et al. 2020).
However, there are still many suitable regions in the Hengduan Mountain Region that have not been surveyed for
Diploderma diversity to date, including the upper Yalong River Valley and its tributaries in southwestern Sichuan
Provinces and sections of Salween River Valley in Eastern Tibet and western Yunnan (Wang et al. 2021a, 2021b).
Furthermore, there are still several species complexes that require detailed taxonomic revisions, particularly the ar-
boreal species of the D. yunnanense complex and D. fasciatum complex in southern China and northern Indochina
such as Myanmar, Thailand, Laos, and Vietnam (Ota 2000; Wang et al. 2017; Liu et al. 2020). Future taxonomic
studies should focus on increasing survey efforts in the Hengduan Mountain Region and resolving the existing taxo-
nomic uncertainties of the recognized species complexes.
For the species along the upper Salween River, distribution ranges of species were under debates. Rao et al.
(2017) first recorded D. laeviventre in the Nujiang Prefecture of Yunnan Province, extending the distribution of the
species southwards over hundreds of kilometers, and such distribution data were accepted by later studies (Sun &
Gao, 2017; Rao 2020). However, Rao’s distribution extension was based on misunderstanding of literature, as point-
ed out by Wang et al. (2019b). Our discovery of D. yangi in the Salween valley next to the provincial border in Tibet
further exclude the possibility of having D. laeviventre in Yunnan, as any population of D. laeviventre in Yunnan
will be allopatric. It is possible that our new species D. yangi extents its distribution southward into Yunnan along
the Salween River, but its distribution would be limited to the north of Qiunatong Village in Gongshan County.
In contrast to the booming taxonomic studies of Diploderma is the lack of the natural history and evolutionary
studies of the genus. While all recent taxonomic studies are of mainland Chinese species, only a few descriptive
ecology and physiology studies have been published on species from the mainland (Li & Li 1981; Li et al. 2004),
and majority of the existing literature on the ecology and evolution of the genus come from a handful of species
from East Asian Islands (Tanaka 1986; Huang 2007; Kuo et al. 2007; Yang et al. 2018). Such lack of knowledge for
most of its diversity on the mainland Asia not only prevents effective conservation of these vulnerable endemic spe-
cies, but it also prevents the discoveries of unique biological findings. Future studies should fill the gap of knowl-
edge and provide scientifically guided conservation plans for these threatened endemic lizards.

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Acknowledgement
We followed IACUC protocols (IACUC R13-11) and relevant protocols of the Animal Care at Southwest Forestry
University for the proper treatments of animals in the field. Collection permit was issued by the Southwest Forestry
University. We thank Mr. Jiansheng Dong for providing photos of the unvouchered individuals of the new species;
Dr. Zhiyong Yuan, Mr. Xiaolong Liu, and Tie Minhua for their assistances in the field; Dr. Veerappan Deepak for
his helps in examining specimens at NHM; and Dr. Jiatang Li (CIB), Mr. Ke Jiang (CIB), Mr. Jinlong Ren (CIB),
Drs. Rafe Brown and Luke Welton (KU), Mr. Wynn Addison (NMNH), Ms. Erica Ely and Ms. Lauren Scheinberg
(CAS), and Drs. James Hanken and Jonathan Losos (MCZ) for their supports in allowing us examine specimens or
facilitate specimen loans.
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APPENDIX I. Examined specimens of recognized species of the genus Diploderma in this study. Museum abbrevia-
tions include the following: Museum of Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ); Museum of
California 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; Univer-
sity of Kansas Biodiversity Institute (KU), Lawrence, KS, USA; Field Museum of Natural History (FMNH), Chicago, IL,
USA; Museum of Comparative Zoology (MCZ), Cambridge, MA, USA; Natural History Museum (NHM), London, UK;
and Smithsonian National Museum of Natural History (USNM).
D. angustelinea (n = 10): KIZ 029703 (holotype), KIZ 044484, 044796, 044797, 029404–029408, 029710 (paratypes), Maid-
ilong Village, Muli, Sichuan, China.
D. aorun (n = 13): KIZ 044735 (holotype), Dari Village, Deqin, Yunnan, China; KIZ 032734, 032736, 032737, 032735, near
Benzilan township, Deqin, Yunnan Province, China; KIZ 044431, 044432, 044433, Rongzong Village, Deqin, Yunnan,
China; KIZ 044740, 044742, near Zhidu, Deqin, Yunnan, China; CIB 116315–16, CIB 116318, from Songmai Township,
Derong, Sichuan China; KIZ 044764, near Derong township, Sichuan, China (all paratypes).
D. bowoense (n = 4): KIZ 044700 (holotype), KIZ 044701, 044703, 044756 (paratypes), from Bowo Village, Muli Tibetan
Autonomous County, Sichuan Province, China.
D. batangense (n = 18): KIZ 019276–019279, 019281, 019282, 019285, 019286, 044729, 044731, 044765, 044766, 044769,
044783, 044787 (topotypes), Batang, Sichuan, China; KIZ 09404, 019312, 019314, Mangkang, Tibet, China.
D. brevicauda (n = 4): KIZ 044497, from the Developing District of Lijiang, 20km northwest from Lijiang City, Yunnan, China;
KIZ 044305, 044306, from the mountain overseeing Lijiang City, Yunnan, China; KIZ 028338, from Shangjiang Village,
Shangrila County, Yunnan, China.

WANG ET AL.
220 · Zootaxa 5099 (2) © 2022 Magnolia Press
D. brevipes (n = 2): CAS 71998 (holotype), CAS 162100, Taiwan, China.
D. chapaense (n = 6): CIB 2679/583623, KIZ 047085, Jingdong, Yunnan Province, China. KIZ 040145, Dali, Yunnan, China;
KIZ 034921–23, Lvchun, Yunnan, China.
D. drukdaypo (n = 8): KIZ 027616 (holotype), KIZ 027618, 027619, 027628–027630 (paratypes), Chaya, Chamdo, Tibet,
China; KIZ 016486 (paratype), Karuo, Chamdo, Tibet, China.
D. dymondi (n = 20): KIZ 040639, 040640 (topotypes), Dongchuan, Yunnan, China; KIZ 040147–040153, CIB 1869/755156,
City of Panzhihua, Panzhihua District, Sichuan, China; KIZ 95I1001, 95I1002, 95I1016, 95I1018, 95I1022, KIZ 040645–
48, Dayao, Yunnan, China; CIB 1870/65I5020, Yuzha, Huili County, Liangshan Prefecture, Sichuan, China.
D. fasciatum (n = 3): CIB 2620–2622, Peng County, Sichuan, China.
D. flaviceps (n = 15): CIB 2234, 2332, 2333, 2341, 2354, 2355, 2549, 2554, 2556, 2561, 2567; KIZ 05181, 05182, 84001 (topo-
types), Luding, Sichuan, China; KU 208076 (topotye), Luding, Sichuan, China.
D. flavilabre (n = 8): KIZ 032693 (holotype), 032692, 032694, 032695–032699, 032730 (paratypes), Yebatan, Gaiyu, Sichuan,
China.
D. formosgulae (n = 15): KIZ 044425 (holotype), 044373, 044417, 044418, 044424, 044427, 044428, 044435, 044375, 044420,
044421, 044423, 044429, 044430, 044437 (paratypes), from Yangla Village, Deqin County, Yunnan Province, China.
D. grahami (n = 1): USMN 65500 (holotype), Yibin, Sichuan, China.
D. hamptoni (n = 1): BMNH 1908.9.18.1/1946.8.14.1 (holotype), Mogok, Myanmar.
D. iadinum (n = 16): KIZ 019321 (holotype), 09398 (allotype), 09401– 03, 019322, 019325–019328 (paratypes), Ninong, De-
qin, Yunnan, China; KIZ 027702–05, Yunling, Deqin, Yunnan, China.
D. laeviventre (n = 5): KIZ 014038 (holotype), 014037, 014041–014043 (paratypes), Markam, Tibet, China.
D. makii (n = 2): MCZ R-172743 (paratype), R-181443, Nantong, Taiwan, China.
D. micangshanense (n = 9): CIB 86348, 86351, Xianyang, Shaanxi, China; CIB 86356, 86357, 86360, 86361, Luonan, Shaanxi,
China; CIB 2572, 2578, 2582, Wenxian, Gansu, China.
D. panchi (n = 4): KIZ 032715 (holotype), 032716, 032717, 032729 (paratypes), Yajiang Township, Ganzi, northwest Sichuan
Province, China.
D. panlong (n = 7): KIZ 040138 (holotype), 040137, 040139, 040140, 040143, 040141, 040142 (paratypes), Miansha Village,
Mianning, Sichuan, China.
D. polygonatum (n = 11): MCZ 45954, 45956 (paratypes), CAS 21215, 21355, 21243, 21221, 21244, Okinawa, Japan.
D. qilin (n = 13): KIZ 028332 (holotype); paratypes: 028333, 028334–336, Balong, Deqin County, Yunnan, China; KIZ 044412,
044413, Baka Village, Shangri-La, Yunnan, China; KIZ 044405, 044407, 044408, Meiding Village, Shangri-La, Yunnan,
China; KIZ 044745, 044744, Cangjue Village, Shangri-La, Yunnan, China; KIZ 044820, Pengnanshou Bridge, Shangri-La,
Yunnan, China.
D. splendidum (n = 6): USNM 35522 (holotype), Yichang, Hubei, China; CIB 2588, 2591, 2596, 72468, 72469, Chongqing,
China.
D. slowinskii (n = 11): KIZ 027541, 027543, 027572–74, 027577, 027579, 027595, 027596, 027598, 027600 (topotypes), Nu-
jiang Prefecture, Yunnan, China.
D. swild (n = 8): KIZ 034912 (holotype), 034894, 034913, 034914, 040125–27 (paratypes), Panzhihua, Sichuan, China; CIB
1871/105074 (paratype), Xichang, Liangshan Prefecture, Sichuan, China.
D. swinhonis (n = 4): CAS 18085, 18089, 18098, 18099, Taiwan, China.
D. varcoae (n = 15): KIZ 85I0006, 85I0009, 83001, 034294 (topotypes), Kunming, Yunnan, China; KIZ 015691, Yuxi, Yun-
nan, China; KIZ 015689, Mengzi, Honghe Prefecture, Yunnan, China; CIB 2682/625219, 2659/625233, 2657/625212,
2660/625213, 2668/625214, 2667/625215, 2666/625217, 2662/625218, 2658/625221, 2679/583623, Jizu Mountain, Dali,
Yunnan Province.
D. vela (n = 31): KIZ 013801 (holotype), KIZ 013802, 013813, 013800, 013805–013811 (paratypes), Jerkalo, Tibet, China;
KIZ 027641, 027645–49, Rumei, Markam, Tibet, China; KIZ 027668, 027670–73, Tongsha, Markam, Tibet, China; KIZ
027667, 027693–695, Foshan, Deqin, Yunnan, China; KIZ 027680, 027681, Xilu, Deqin, Yunnan, China; KIZ 027682,
027684, Xidang, Deqin, Yunnan, China.
D. yulongense (n = 15): KIZ 09399, 09400, 028291–028293, 028294, 028296–028298, 028299, 028300, 028303, 028342–
028344, Xianggelila County, Yunnan, China.
D. yunnanense (n = 8): CIB 2684, 2686, 2687, 2689, KIZ 82081 (topotypes), Longling, Yunnan, China; KIZ 74II0240, 0248,
79I469, Tengchong, Yunnan, China; KIZ 040193, Yingjiang, Yunnan, China.
D. zhaoermii (n = 14): CIB 2690 (holotype), 86432, 86435, 85721, 85722, 86433, 86434, 86436, Wenchuan, Sichuan, China;