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Research Article
Four new species of the genus Diploderma Hallowell, 1861 (Squamata, Agamidae) from China
expand article infoShuo Liu‡§, Mian Hou|, Natalia B. Ananjeva, Dingqi Rao§
‡ Kunming Natural History Museum of Zoology, Kunming, China
§ Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China
| Sichuan Normal University, Chengdu, China
¶ Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia

Corresponding author: Natalia B. Ananjeva ( natalia.ananjeva@zin.ru )

Corresponding author: Dingqi Rao ( raodq@mail.kiz.ac.cn )

Academic editor: Thomas Ziegler
© 2023 Shuo Liu, Mian Hou, Natalia B. Ananjeva, Dingqi Rao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Liu S, Hou M, Ananjeva NB, Rao D (2023) Four new species of the genus Diploderma Hallowell, 1861 (Squamata, Agamidae) from China. ZooKeys 1148: 167-207. https://doi.org/10.3897/zookeys.1148.97706
ZooBank: urn:lsid:zoobank.org:pub:80D02AB5-0175-4054-AF53-67FAB0654750
Open Access

Abstract

Four new species of Diploderma are described from Sichuan and Yunnan provinces, southwestern China, based on an integrative taxonomic approach, combining morphological and genetic data. The first new species from Danba County, Sichuan Province, is morphologically most similar and phylogenetically closely related to D. flaviceps, but it can be diagnosed from the latter by having a relatively much shorter tail and by a genetic distance of 4.4% in the ND2 gene; the second new species from Muli County, Sichuan Province, is phylogenetically closely related to D. daochengense, D. yongshengense, and D. yulongense, but it can be diagnosed from the latter three species by having a pale yellow gular spot and by genetic distances of 5.6–6.7% in the ND2 gene; the third new species from Jiulong County, Sichuan Province, is morphologically most similar and phylogenetically closely related to D. angustelinea, but it can be diagnosed from the latter by having a relatively much longer tail and by a genetic distance of 2.8% in the ND2 gene; and the last new species from Weixi County, Yunnan Province, is phylogenetically closely related to D. aorun, but it can be diagnosed from the latter by having a pale yellow gular spot and by a genetic distance of 2.9% in the ND2 gene. Our work brings the number of species within the genus Diploderma to 46.

Keywords

Hengduan Mountain Region, ND2, Sichuan, systematic, taxonomy, Yunnan

Introduction

Currently the genus Diploderma Hallowell, 1861 comprises 42 recognized species (Cai et al. 2022; Liu et al. 2022; Wang et al. 2022b) which are mainly distributed in the hot-dry river valleys of the Hengduan Mountain Region of southwestern China, and more than half of them were described in the past decade (Wang et al. 2021a, 2022a; Cai et al. 2022; Liu et al. 2022; Uetz et al. 2022).

The Hengduan Mountain Region contains many high mountains and rivers, forming complex hot-dry river valleys in this region, which are suitable for Diploderma species to inhabit. Therefore, it is not surprising that numerous new species of Diploderma were found in this region. However, due to the complex terrain and inconvenient transportation, there are still many areas of this region that have not been fully investigated for biodiversity.

During our field surveys in Yunnan and Sichuan provinces, China, in 2022, some specimens of Diploderma were collected from four different sites in the Hengduan Mountain Region. In terms of morphology, these specimens showed distinct differences to any recognized species of the genus. Furthermore, phylogenetic analyses indicated differentiation of these populations also at a molecular level. By comparing our data with morphological and molecular differentiation within existing species of Diploderma, we conclude that these populations have exceeded an intraspecific variation level and thus describe four new species of Diploderma herein.

Materials and methods

Sampling

Field survey in Yunnan Province was conducted from June to July 2022 by Shuo Liu, and field survey in Sichuan Province was conducted from July to August 2022 by Mian Hou. Specimens were collected in the morning or afternoon from Weixi County, Yunnan Province, and Danba, Muli, and Jiulong counties, Sichuan Province, respectively (Fig. 1). Photographs were taken to document colour patterns in life prior to anaesthetisation and euthanasia, respectively. Genetic tissues were stored in 99% ethanol taken from livers, and specimens were preserved in 75% ethanol. All newly collected specimens were deposited at Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ).

Figure 1. 

Map showing the type localities of Diploderma danbaense sp. nov. in Danba County, Ganzi Prefecture, Sichuan Province (black dot), Diploderma donglangense sp. nov. in Muli County, Liangshan Prefecture, Sichuan Province (black pentagon), Diploderma jiulongense sp. nov. in Jiulong County, Ganzi Prefecture, Sichuan Province (black triangle), and Diploderma tachengense sp. nov. in Weixi County, Diqing Prefecture, Yunnan Province (black square), in the Hengduan Mountain Region, southwestern China. The elevation data were obtained from Geospatial Data Cloud (2022).

Morphological characteristics and analyses

Morphometric measurements and pholidosis data were taken for all newly collected specimens and 27 museum specimens deposited at KIZ. Specimens were measured using a digital caliper to the nearest 0.1 mm. Measurements were taken on the left side (unless the left side was damaged, then the right side was used), values for paired pholidosis characteristics were recorded on both sides and were provided in left/right order (following Liu et al. 2020, 2022; Wang et al. 2022a, b). The following morphometric characteristics were measured and counted following Liu et al. (2022) and Wang et al. (2022a):

F4S subdigital lamellae under fourth finger, subdigital lamellae scales from the base between third and fourth finger to the tip of fourth finger, excluding the claw;

FLL fore-limb length, measured between the point of insertion at axillary to the tip of fourth finger, excluding the claw, measured on straightened limb;

HD head depth, measured as the perpendicular distance at the temporal region of head;

HL head length, measured from the tip of snout to the rear border of the angle of jaw;

HLL hind limb length, measured between the point of insertion at groin to the tip of fourth toe, excluding the claw, measured on straightened limb;

HW head width, measured between the widest points of the head;

IL infralabial scale number, enlarged, modified labial scales from mental to the corner of mouth;

MD middorsal crest scale number, modified crest scales longitudinally from the first nuchal crest to the scale above cloaca;

NSL nasal–supralabials scale rows, number of horizontal rows of small scales between the first supralabial and the nasal;

SEL snout–eye length, measured between the tip of snout and anterior edge of orbital bone;

SL supralabial scale number, enlarged, modified labial scales from rostral to the corner of mouth;

SOR suborbital scale rows, longitudinal rows of scales between supralabials and inferior-most edge of orbit circle, excluding fine ciliary scales in the orbit;

SVL snout–vent length, measured from the snout tip to anterior edge of the cloaca;

T4L fourth toe length, measured between the tip of fourth toe to the base between third and fourth toe, excluding the claw;

T4S subdigital lamellae under fourth toe, subdigital lamellae scales from the base between third and fourth toe to the tip of fourth toe, excluding the claw;

TAL tail length, measured from the anterior edge of the cloaca to the tip of tail;

TRL trunk length, measured between the limb insertion points between axillary and groin;

VN ventral scale number, ventral body scales counted in a straight line along the medial axis between the transverse gular fold and the anterior edge of cloaca.

We compared morphological characters of the new species with other members of the genus relying on examined specimens and original species descriptions (Hallowell 1861; Günther 1864; Anderson 1878; Boulenger 1906, 1918; Barbour and Dunn 1919; Stejneger 1924; Mertens 1926; Smith 1935; Gressitt 1936; Bourret 1937; Song 1987; Ota 1989; Ota et al. 1998; Li et al. 2001; Gao and Hou 2002; Manthey et al. 2012; Wang et al. 2015, 2016, 2019b, d, 2021a, b, 2022a; Ananjeva et al. 2017; Rao et al. 2017; Liu et al. 2020, 2022; Cai et al. 2022), and the additional data from Wu et al. (2005), Manthey (2008), and Wang et al. (2017, 2018, 2019b, c, 2021a, 2022b).

Principal component analysis (PCA) was used to determine whether the newly collected specimens and species with closely morphological similarities to them occupied unique positions in morphospace, and whether the results coincide with the delineation of species as indicated by molecular phylogenetic analysis. Characters used in the PCA were mensural data from SVL, TAL, HL, HW, HD, SEL, FLL, HLL, TRL, and T4L. PCA was performed using the prcomp command in R 4.2.2 (R Core Team 2022). The scatter plot was plotted using the ggplot2 package in R 4.2.2. As species of Diploderma are known to be sexually dimorphic in morphometric measurements (Manthey et al. 2012; Wang et al. 2015, 2018, 2019b), mensural data of each sex are treated separately for analyses.

Molecular analysis

Total genomic DNA for the newly collected specimens was extracted from liver tissues with a standard three step phenol chloroform extraction method (Sambrook et al. 1989). The mitochondrial gene NADH dehydrogenase subunit 2 (ND2) was amplified and sequenced by using published primers (Wang et al. 2019a). PCR and sequencing methods were the same as Liu et al. (2020). All new sequences were edited and manually managed using SeqMan in Lasergene 7.1 (DNASTAR Inc., Madison, WI, USA) and MEGA 11 (Tamura et al. 2021). Representative species of Pseudocalotes Fitzinger were chosen as outgroups (Liu et al. 2022). Genetic data for 39 species of Diploderma and two species of outgroup taxa were obtained from GenBank (Table 1). The technical computation methods for sequences alignment, best substitution model selection, Bayesian inference (BI) and maximum likelihood (ML) phylogenetic analyses, and genetic divergences calculation were the same as those in Liu et al. (2022).

Table 1.
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GenBank accession numbers for the sequences used in this study.

Species Voucher Locality Accession
Diploderma angustelinea KIZ 029704 Muli, Sichuan, China MT577930
KIZ 029705 Muli, Sichuan, China MT577924
KIZ 029708 Muli, Sichuan, China MT577931
KIZ 029710 Muli, Sichuan, China MT577927
KIZ 032488 Muli, Sichuan, China MT577925
KIZ 032489 Muli, Sichuan, China MT577926
KIZ 032490 Muli, Sichuan, China MT577928
KIZ 032491 Muli, Sichuan, China MT577929
Diploderma aorun KIZ 032733 Benzilan, Yunnan, China MT577938
KIZ 032734 Benzilan, Yunnan, China MT577939
KIZ 032735 Benzilan, Yunnan, China MT577937
KIZ 032736 Benzilan, Yunnan, China MT577936
KIZ 032737 Benzilan, Yunnan, China MT577940
Diploderma batangense KIZ 09404 Zhubalong, Tibet, China MK001412
KIZ 019276 Batang, Sichuan, China MK001413
Diploderma brevicauda KIZ 044305 Lijiang, Yunnan, China MW506021
KIZ 044306 Lijiang, Yunnan, China MW506022
Diploderma bowoense KIZ 044757 Muli, Sichuan, China MW506020
KIZ 044758 Muli, Sichuan, China MW506019
Diploderma brevipes NMNS 19607 Taiwan, China MK001429
NMNS 19608 Taiwan, China MK001430
Diploderma chapaense KIZ 034923 Lvchun, Yunnan, China MG214263
ZMMU NAP-01911 Chapa, Vietnam MG214262
Diploderma daochengense 20210905 Muli, Sichuan, China OP595620
DC001 Daocheng, Sichuan, China OP595621
DC003 Daocheng, Sichuan, China OP595622
DC004 Daocheng, Sichuan, China OP595623
20210904 Muli, Sichuan, China OP595624
Diploderma drukdaypo KIZ 027627 Jinduo, Tibet, China MT577950
KIZ 027628 Zhuka, Tibet, China MT577952
Diploderma dymondi KIZ 040639 Dongchuan, Yunnan, China MK001422
KIZ 040640 Dongchuan, Yunnan, China MK001423
Diploderma fasciatum SYS r002175 Wuming, Guangxi, China OM055809
KIZ 040192 Daweishan, Yunnan, China OM055800
Diploderma flaviceps KIZ 01851 Luding, Sichuan, China MK001416
KIZ 01852 Luding, Sichuan, China MK001417
KIZ 019575 Kangding, Sichuan, China MT577896
KIZ 019576 Kangding, Sichuan, China MT577897
KIZ 019577 Kangding, Sichuan, China MT577895
KIZ 019578 Kangding, Sichuan, China MT577894
KIZ 019579 Kangding, Sichuan, China MT577898
Diploderma flavilabre KIZ 032692 Baiyu,Sichuan, China MT577916
KIZ 032694 Baiyu,Sichuan, China MT577917
Diploderma formosgulae KIZ 044420 Deqin, Yunnan, China MW506024
KIZ 044421 Deqin, Yunnan, China MW506025
Diploderma iadinum KIZ 027697 Yunling, Yunnan, China MT577956
KIZ 027702 Yunling, Yunnan, China MT577957
Diploderma kangdingense 20210916 Kangding, Sichuan, China OP595625
20210917 Kangding, Sichuan, China OP595626
Diploderma laeviventre KIZ 014037 Basu, Tibet, China MK001407
KIZ 027691 Basu, Tibet, China MT577892
Diploderma limingense KIZ2022015 Yulong, Yunnan, China OP428783
KIZ2022017 Yulong, Yunnan, China OP428784
Diploderma luei NMNS 19604 Taiwan, China MK001433
NMNS 19605 Taiwan, China MK001434
Diploderma makii NMNS 19609 Taiwan, China MK001431
NMNH 19610 Taiwan, China MK001432
Diploderma menghaiense KIZ L0030 Menghai, Yunnan, China MT598655
KIZ L0031 Menghai, Yunnan, China MT598656
Diploderma micangshanense KIZ 032801 Shiyan, Hubei, China MK578665
KIZ 023231 Xixia, Henan, China MK578664
Diploderma panchi KIZ 032715 Yajiang, Sichuan, China MT577946
KIZ 032716 Yajiang, Sichuan, China MT577944
Diploderma panlong KIZ 040137 Miansha, Sichuan, China MT577906
KIZ 040138 Miansha, Sichuan, China MT577907
Diploderma polygonatum NMNS 19598 Taiwan, China MK001427
NMNS 19599 Taiwan, China MK001428
Diploderma qilin KIZ 028332 Deqin, Yunnan, China MT577941
KIZ 028333 Deqin, Yunnan, China MT577942
Diploderma shuoquense KIZ2022004 Xiangcheng, Sichuan, China OP428773
KIZ2022005 Xiangcheng, Sichuan, China OP428774
Diploderma slowinskii CAS 214906 Gongshan, Yunnan, China MK001405
CAS 214954 Gongshan, Yunnan, China MK001406
Diploderma splendidum KIZ 015973 Yichang, Hubei, China MK001418
LSUMZ 81212 Unknown AF288230
Diploderma swild KIZ 034914 Panzhihua, Sichuan, China MN266299
KIZ 034894 Panzhihua, Sichuan, China MN266300
Diploderma swinhonis NMNS 19592 Taiwan, China MK001419
NMNS 19593 Taiwan, China MK001420
Diploderma varcoe WK-JK 011 Yuxi, Yunnan, China MT577903
KIZ 026132 Mengzi, Yunnan, China MK001421
Diploderma vela KIZ 019299 Quzika, Tibet, China MK001414
KIZ 034925 Quzika, Tibet, China MK001415
Diploderma xinlongense 20210907 Xinlong, Sichuan, China OP595613
20210908 Xinlong, Sichuan, China OP595614
Diploderma yangi SWFU 005410 Chayu, Tibet, China OL449603
SWFU 005412 Chayu, Tibet, China OL449604
Diploderma yongshengense KIZ2022009 Yongsheng, Yunnan, China OP428777
KIZ2022008 Yongsheng, Yunnan, China OP428778
KIZ2022010 Yongsheng, Yunnan, China OP428779
KIZ2022011 Yongsheng, Yunnan, China OP428780
Diploderma yulongense KIZ 028291 Hutiaoxia, Yunnan, China MT577921
KIZ 028292 Hutiaoxia, Yunnan, China MT577922
KIZ 028300 Baishuitai, Yunnan, China MT577923
KIZ 09399 Xianggelila, Yunnan, China MK001410
KIZ 043196 Xianggelila, Yunnan, China MK001411
Diploderma yunnanense CAS 242271 Baoshan, Yunnan, China MK001408
KIZ 040193 Yingjiang, Yunnan, China MK578658
Diploderma zhaoermii KIZ 019564 Wenchuan, Sichuan, China MK001425
KIZ 019565 Wenchuan, Sichuan, China MK001426
Diploderma danbaense sp. nov. KIZ2022048 Danba, Sichuan, China OQ378180
KIZ2022049 Danba, Sichuan, China OQ378181
KIZ2022050 Danba, Sichuan, China OQ378182
KIZ2022051 Danba, Sichuan, China OQ378183
KIZ2022056 Danba, Sichuan, China OQ378184
Diploderma donglangense sp. nov. KIZ2022057 Muli, Sichuan, China OQ378185
KIZ2022058 Muli, Sichuan, China OQ378186
KIZ2022059 Muli, Sichuan, China OQ378187
KIZ2022060 Muli, Sichuan, China OQ378188
KIZ2022061 Muli, Sichuan, China OQ378189
Diploderma jiulongense sp. nov. KIZ2022086 Jiulong, Sichuan, China OQ378190
KIZ2022087 Jiulong, Sichuan, China OQ378191
KIZ2022099 Jiulong, Sichuan, China OQ378192
KIZ2022100 Jiulong, Sichuan, China OQ378193
KIZ2022101 Jiulong, Sichuan, China OQ378194
Diploderma tachengense sp. nov. KIZ2022028 Weixi, Yunnan, China OQ378195
KIZ2022027 Weixi, Yunnan, China OQ378196
KIZ2022029 Weixi, Yunnan, China OQ378197
KIZ2022038 Weixi, Yunnan, China OQ378198
KIZ2022039 Weixi, Yunnan, China OQ378199
Pseudocalotes brevipes MVZ 224106 Vinh Phuc, Vietnam AF128502
Pseudocalotes kakhiensis KIZ 015975 Gongshan, Yunnan, China MK001435

Results

The obtained sequence alignment is 1028 bp in length. The resulting topologies from BI and ML analyses are consistent (Fig. 2). The specimens from Danba County formed a clade sister to Diploderma flaviceps (Barbour & Dunn, 1919) with strong support; the specimens from Muli County formed a clade sister to the clade consisting of D. daochengense Cai, Zhang, Li, Du, Xie, Hou, Zhou & Jiang, 2022, D. yongshengense Liu, Hou, Rao & Ananjeva, 2022, and D. yulongense (Manthey, Denzer, Hou & Wang, 2012) with strong support; the specimens from Jiulong County formed a clade sister to D. angustelinea Wang, Ren, Wu, Che & Siler, 2020 with strong support; and the specimens from Weixi County formed a clade sister to D. aorun Wang, Jiang, Zheng, Xie, Che & Siler, 2020 with strong support. The minimum uncorrected pairwise distance between the specimens from Danba County and other species of Diploderma is 4.4% (between the population from Danba County and D. flaviceps), the minimum uncorrected pairwise distance between the specimens from Muli County and other species of Diploderma is 5.6% (between the population from Muli County and D. yulongense), the minimum uncorrected pairwise distance between the specimens from Jiulong County and other species of Diploderma is 2.8% (between the population from Jiulong County and D. angustelinea), and the minimum uncorrected pairwise distance between the specimens from Weixi County and other species of Diploderma is 2.9% (between the population from Weixi County and D. aorun), which are all greater than the genetic distance (2.6%) between the two recognized species D. drukdaypo and D. vela (Suppl. material 1).

Figure 2. 

Bayesian phylogram of the genus Diploderma inferred from mitochondrial gene ND2 (1028 bp). Numbers before slashes indicate Bayesian posterior probabilities and numbers after slashes indicate ML ultrafast bootstrap values. The symbol “–” represents the value below 0.90/90.

The first two major principal components from PCA were retained, which accounted for 83.32%–91.12% of the total variances. The scatter plot based on PC1 and PC2 showed that the newly collected specimens and species with closely morphological similarities to them can be segregated in both sexes (Fig. 3).

Figure 3. 

PCA based on ten morphometric characteristics (SVL, TAL, HL, HW, HD, SEL, FLL, HLL, TRL, and T4L) for males of Diploderma flaviceps and Diploderma danbaense sp. nov. (upper left), females of D. flaviceps and Diploderma danbaense sp. nov. (upper right), males of D. angustelinea and Diploderma jiulongense sp. nov. (lower left), and females of D. angustelinea and Diploderma jiulongense sp. nov. (lower right).

Taxonomy

Diploderma danbaense sp. nov.

https://zoobank.org/08AA2744-059D-4295-AAEC-617A595FC3C0
Figs 4, 5, 6

Type material

Holotype. KIZ2022048, adult male, collected on 6 August 2022 by Mian Hou from Bawang Township, Danba County, Ganzi Prefecture, Sichuan Province, China (30°58'59"N, 101°52'29"E, 2020 m elevation).

Paratypes. KIZ2022049, KIZ2022050, KIZ2022056, three adult males; KIZ2022051, one adult female; collecting information the same as the holotype.

Etymology

The specific epithet refers to Danba County, where the new species was discovered.

Diagnosis

Diploderma danbaense sp. nov. can be diagnosed from congeners by a combination of the following morphological characteristics: (1) body size large, SVL 68.7–77.0 mm in adult males, 76.6 in adult female; (2) tail short, TAL/SVL 1.61–1.78 in adult males, 1.55 in adult female; (3) head relatively long, HW/HL 0.66–0.75 in adult males, 0.63 in adult female; (4) limbs moderately long, FLL/SVL 0.41–0.47 in adult males, 0.44 in adult female, HLL/SVL 0.66–0.70 in adult males, 0.65 in adult female; (5) MD 48–58; (6) F4S 16–20, T4S 21–26; (7) tympanum concealed; (8) nuchal and dorsal crests discontinuous, scales of nuchal and dorsal crests enlarged, strongly erected skin fold under nuchal crest and moderately erected skin fold under dorsal crest in males in life, weakly erected skin fold under nuchal crest and no skin fold under dorsal crest in females in life; (9) distinct transverse gular fold present; (10) ventral scales of head heterogeneous in size, anterior and middle ones larger, posterior and side ones smaller, all strongly keeled; (11) ventral scales of body strongly keeled; (12) gular spot absent in both sexes; (13) dorsolateral stripes distinct in males, strongly jagged, pale yellow in life; (14) a series of dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum; (15) a distinct wide black stripe on shoulder fold region on each side; (16) stripes around eye absent or very indistinct; (17) oral cavity, inner lips, and tongue pale flesh colour in life.

Description of holotype

Adult male, SVL 77.0 mm; tail short, TAL 130.0 mm, TAL/SVL 1.69; limbs moderately long, FLL 33.1 mm on left side, FLL/SVL 0.43, HLL 53.5 mm on left side, HLL/SVL 0.69. Head relatively long, HW/HL 0.67, HD/HW 0.84; snout moderately long, SEL/HL 0.37. Rostral elongated, bordered by five small postrostral scales; dorsal head scales heterogeneous, all strongly keeled; distinct Y-shaped ridge on dorsal snout. Nasal oval, separated from first supralabial by two rows of scales on each side; loreals small, keeled; suborbital scale rows 5/4, keeled; canthus rostralis elongated, scales greatly overlapping each other; enlarged, keeled scales forming single lateral ridge from posteroinferior eye to posterosuperior tympanum on each side; tympanum concealed under scales; SL 10/10, smooth. Mental pentagonal; IL 11/11; enlarged chin shields 10/10, smooth, first one connected IL on each side, second one separated from IL by one row of small scales on each side, remaining ones separated from IL by two rows of small scales on each side; ventral head scales heterogeneous in size, anterior and middle ones larger, posterior and lateral ones smaller, all strongly keeled; distinct transverse gular fold present; gular pouch moderately developed.

Distinct shoulder fold present; dorsal body scales heterogeneous in size and shape, all keeled, tip pointing backwards; axillary scales much smaller than remaining dorsals; enlarged dorsal scales forming one intermittent longitudinal row between dorsal crest and dorsolateral stripe on each side, remaining enlarged dorsal scales irregularly scattered on each side of body. Nuchal and dorsal crests discontinuous, separated by a diastema, scales of nuchal and dorsal crests enlarged; strongly erected skin fold under nuchal crest and moderately erected skin fold under dorsal crest; MD 48. Dorsal limb scales strongly keeled, homogeneous on fore-limbs and heterogeneous on hind limbs; F4S 17/16, T4S 21/22. Ventral body scales approximately parallel, homogeneous, all strongly keeled, VN 65. Ventral limb scales parallel, small on upper arms and thighs and larger on forearms and crus, all strongly keeled. Tail scales all strongly keeled, ventral tail scales slightly larger than dorsal tail scales.

Colouration of holotype in life

Dorsal surface of head brownish grey. Two indistinct black transverse bands between orbits on dorsal surface of head. Lateral surfaces of head grey. Two indistinct black stripes from posteroinferior eye to anterior tympanum on each side. Upper lips grey, lower lips white. Oral cavity, inner lips, and tongue with pale flesh colour.

Dorsal surface of body purplish grey. A pale yellow strongly jagged dorsolateral stripe from neck to pelvis on each side of body. A series of dark, hollow, approximately rhomboid-shaped patterns between dorsolateral stripes from neck to base of tail, hollow region pale yellow. A distinct wide black stripe on shoulder fold region on each side. Some irregular black patches below dorsolateral stripe on each side of body, no pale spots on each side of body. Dorsal surfaces of limbs brownish grey with dark transverse bands. Dorsal surface of tail greyish white with distinct dark transverse bands.

Ventral surface of head white with distinct grey reticulated pattern. No gular spot. Ventral surfaces of body and limbs greyish white with no pattern, ventral surface of tail greyish white with indistinct dark transverse bands.

Variations

The variations of metrical characteristics of the type series are provided in Table 2. Other variations are as follows: the transverse bands on the dorsal surface of head and the stripes posteroinferior to the eye are more indistinct in all paratypes; the skin fold under nuchal crest is more weak, no skin fold under dorsal crest, and the dorsolateral stripes are indistinct, pale grey in the female paratype.

Table 2.
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Morphological data of the type series of Diploderma danbaense sp. nov. Morphometric measurements are in the unit of mm. For measurement and count methods and abbreviations, see the Materials and methods.

KIZ2022048 holotype ♂ KIZ2022049 paratype ♂ KIZ2022050 paratype ♂ KIZ2022051 paratype ♀ KIZ2022056 paratype ♂
SVL 77.0 70.0 68.7 76.6 69.8
TAL 130.0 116.6 122.6 119.1 112.7
HL 26.6 22.7 22.6 23.5 21.4
HW 17.9 16.8 15.0 14.7 16.1
HD 15.0 14.1 12.8 12.8 12.9
SEL 9.9 8.2 8.2 8.4 7.5
FLL 33.1 32.8 32.6 33.6 28.7
HLL 53.5 49.3 48.4 50.0 46.4
T4L 12.2 12.0 12.4 12.1 11.9
TRL 33.2 31.3 29.2 34.5 31.8
TAL/SVL 1.69 1.67 1.78 1.55 1.61
SEL/HL 0.37 0.36 0.36 0.36 0.35
HW/HL 0.67 0.74 0.66 0.63 0.75
HD/HW 0.84 0.84 0.85 0.87 0.80
FLL/SVL 0.43 0.47 0.47 0.44 0.41
HLL/SVL 0.69 0.70 0.70 0.65 0.66
TRL/SVL 0.43 0.45 0.43 0.45 0.46
SL 10/10 9/9 10/9 11/10 10/10
IL 11/11 11/11 11/11 11/11 10/10
NSL 2/2 2/1 1/1 2/2 2/2
MD 48 50 47 58 53
F4S 17/16 16/17 18/18 20/19 19/20
T4S 21/22 22/23 24/23 24/26 25/24
SOR 5/4 4/4 4/5 5/4 5/5
VN 65 72 66 74 68

Comparisons

Diploderma danbaense sp. nov. differs from D. brevipes (Gressitt, 1936), D. chapaense (Bourret, 1937), D. fasciatum (Mertens, 1926), D. hamptoni (Smith, 1935), D. luei (Ota, Chen & Shang, 1998), D. makii (Ota, 1989), D. menghaiense Liu, Hou, Wang, Ananjeva & Rao, 2020, D. micangshanense (Song, 1987), D. ngoclinense (Ananjeva, Orlov & Nguyen, 2017), D. polygonatum Hallowell, 1861, D. swinhonis (Günther, 1864), and D. yunnanense (Anderson, 1878) by the presence of a transverse gular fold (vs. absence).

Diploderma danbaense sp. nov. differs from D. dymondi (Boulenger, 1906), D. panlong Wang, Che & Siler, 2020, D. slowinskii (Rao, Vindum, Ma, Fu & Wilkinson, 2017), D. varcoae (Boulenger, 1918), and D. swild Wang, Wu, Jiang, Chen, Miao, Siler & Che, 2019 by having concealed tympana (vs. exposed).

Diploderma danbaense sp. nov. differs from D. angustelinea, D. aorun, D. batangense (Li, Deng, Wu & Wang, 2001), D. bowoense Wang, Gao, Wu, Siler & Che, 2021, D. brevicauda (Manthey, Denzer, Hou & Wang, 2012), D. daochengense, D. flavilabre Wang, Che & Siler, 2020, D. formosgulae Wang, Gao, Wu, Dong, Shi, Qi, Siler & Che, 2021, D. iadinum (Wang, Jiang, Siler & Che, 2016), D. laeviventre (Wang, Jiang, Siler & Che, 2016), D. limingensis Liu, Hou, Rao & Ananjeva, 2022, D. qilin Wang, Ren, Che & Siler, 2020, D. xinlongense Cai, Zhang, Li, Du, Xie, Hou, Zhou & Jiang, 2022, D. yangi Wang, Zhang & Li, 2022, D. yongshengense, D. yulongense, and D. zhaoermii (Gao & Hou, 2002) by the absence of a gular spot in males in life (vs. presence of a colourful gular spot).

Diploderma danbaense sp. nov. differs from D. drukdaypo (Wang, Ren, Jiang, Zou, Wu, Che & Siler, 2019) by having strongly keeled ventral scales of body (vs. smooth or weakly keeled); from D. grahami (Stejneger, 1924) by having relatively longer hind limbs (HLL/SVL 0.65–0.70 vs. 0.61), having a distinct transverse gular fold (vs. feeble), the absence of a gular spot after preservation (vs. presence), and the presence of dorsolateral stripes (vs. absence); from D. kangdingense Cai, Zhang, Li, Du, Xie, Hou, Zhou & Jiang, 2022 by the presence of distinct, dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum (vs. absence) and having greyish white ventrolateral surface of body in males in life (vs. yellow); from D. panchi Wang, Zheng, Xie, Che & Siler, 2020 by the presence of distinct, dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum (vs. absence) and the presence of a skin fold under nuchal crest in females in life (vs. absence); from D. shuoquense Liu, Hou, Rao & Ananjeva, 2022 by having strongly keeled ventral head scales (vs. smooth or weakly keeled), the absence of distinct radial stripes around the eyes (vs. presence), and the presence of skin folds under nuchal and dorsal crests in males in life (vs. absence); from D. splendidum (Barbour & Dunn, 1919) by having strongly jagged dorsolateral stripes in males (vs. smooth); and from D. vela (Wang, Jiang & Che, 2015) by having discontinuous nuchal and dorsal crests and skin folds in males in life (vs. continuous nuchal and dorsal crests on continuous skin fold).

Diploderma danbaense sp. nov. is phylogenetically sister to and most similar in morphology characteristic and colouration to D. flaviceps; however, Diploderma danbaense sp. nov. can be differentiated from the latter by having a relatively shorter tail (TAL/SVL 1.61–1.78 vs. 1.88–2.09 in males, 1.55 vs. 1.73–2.17 in females), having relatively shorter hind limbs (HLL/SVL 0.66–0.70 vs. 0.72–0.80 in males, 0.65 vs. 0.70–0.81 in females), having a smaller ratio of head width to head length (HW/HL 0.66–0.75 vs. 0.76–0.84 in males, 0.63 vs. 0.71–0.78 in females), having a greater ratio of head depth to head width (HD/HW 0.80–0.85 vs. 0.70–0.78 in males, 0.87 vs. 0.75–0.83 in females), having relatively shorter fourth toes (T4L/SVL 0.16–0.18 vs. 0.18–0.21 in males, 0.16 vs. 0.17–0.21 in females), and having a moderately erected skin fold under dorsal crest in males in life (vs. strongly erected) and the absence of a skin fold under dorsal crest in females in life (vs. presence).

Figure 4. 

Dorsal view (upper) and ventral view (lower) of the type series of Diploderma danbaense sp. nov. in preservative.

Figure 5. 

Holotype (KIZ2022048) of Diploderma danbaense sp. nov. in life A dorsal view B lateral view C ventral view D close-up view of the dorsolateral side of the head E close-up view of the ventral side of the head F close-up view of the oral cavity.

Figure 6. 

Paratypes of Diploderma danbaense sp. nov. in life A, B the male paratype KIZ2022049 C, D the male paratype KIZ2022050 E, F the female paratype KIZ2022051

Distribution

This species is currently known only from its type locality in Danba County, Ganzi Prefecture, Sichuan Province, China (Fig. 1).

Natural history

This species is terrestrial, inhabiting the hot-dry valley of the upper Dadu River. There are a few trees and many rocks at the type locality (Fig. 16A, B). All specimens were collected between 11 and 12 a.m. when they were basking on rock piles.

Diploderma donglangense sp. nov.

https://zoobank.org/D964D8B1-6004-457A-95E0-0B6EAA97B7E0
Figs 7, 8, 9

Type material

Holotype. KIZ2022057, adult male, collected on 31 July 2022 by Mian Hou from Yaying Village, Donglang Township, Muli County, Liangshan Prefecture, Sichuan Province, China (28°48'49"N, 100°35'42"E, 2750 m elevation).

Paratypes. KIZ2022058–KIZ2022059, KIZ2022061, three adult females; KIZ2022060, adult male; collecting information all the same as the holotype.

Etymology

The specific epithet refers to Donglang Township, where the new species was discovered.

Diagnosis

Diploderma donglangense sp. nov. can be diagnosed from congeners by a combination of the following morphological characteristics: (1) body size relatively small, SVL 44.9–52.8 mm in adult males, 47.2–56.4 in adult females; (2) tail relatively short, TAL/SVL 1.76–1.87 in adult males, 1.59–1.89 in adult females; (3) limbs moderately long, FLL/SVL 0.46–0.47 in adult males, 0.44–0.50 in adult females, HLL/SVL 0.73–0.75 in adult males, 0.70–0.78 in adult females; (4) head moderately long, HW/HL 0.69–0.71 in adult males, 0.71–0.73 in adult females; (5) MD 37–43; (6) F4S 13–17, T4S 19–24; (7) tympanum concealed; (8) nuchal and dorsal crest scales feebly developed, no skin fold under nuchal and dorsal crests; (9) distinct transverse gular fold present; (10) ventral scales of head homogeneous in size, keeled; (11) ventral scales of body strongly keeled; (12) gular spot present in males, present or absent in females, pale yellow in life; (13) dorsolateral stripes distinct in males, moderately jagged, creamy yellow in life; (14) radial stripes around eye indistinct; (15) oral cavity, inner lips, and tongue pale pink in life.

Description of holotype

Adult male, SVL 52.8 mm; tail relatively short, TAL 93.1 mm, TAL/SVL 1.76; limbs moderately long, FLL 24.4 mm on left side, FLL/SVL 0.46, HLL 38.8 mm on left side, HLL/SVL 0.73. Head moderately long, HW/HL 0.69, HD/HW 0.79; snout relatively short, SEL/HL 0.33. Rostral rectangular, bordered by six small postrostral scales; dorsal head scales heterogeneous, all strongly keeled; distinct Y-shaped ridge on dorsal snout. Nasal oval, separated from first supralabial by two rows of scales on each side; loreals small, keeled; suborbital scale rows 5/4, keeled; canthus rostralis elongated, greatly overlapping with each other; enlarged, keeled scales forming distinct single lateral ridge from posteroinferior eye to posterosuperior tympanum on each side; tympanum concealed under scales; SL 10/9, feebly keeled. Mental pentagonal; IL 11/11; enlarged chin shields 4/4, smooth, first one contacting IL on each side, remaining ones separated from IL by one or two rows of small scales on each side; ventral head scales homogeneous in size, keeled; distinct transverse gular fold present; gular pouch weakly developed.

Distinct shoulder fold present; dorsal body scales heterogeneous in size and shape, all keeled, tip pointing backwards; axillary scales much smaller than remaining dorsals; enlarged dorsal scales forming one continuous longitudinal row between dorsal crest and dorsolateral stripe on each side, remaining enlarged dorsal scales roughly forming three or four longitudinal rows on each side of body. Nuchal and dorsal crests feebly developed; no skin fold under nuchal and dorsal crests; MD 40. Dorsal limb scales strongly keeled, mostly homogeneous, except a few enlarged, conical scales on postaxial thighs; F4S 16/16, T4S 24/23. Ventral body scales approximately parallel, homogeneous, all strongly keeled, VN 63. Ventral limb scales parallel, homogeneous, approximately equal in size to ventrals, all strongly keeled. Tail scales all strongly keeled, ventral tail scales larger than dorsal tail scales.

Colouration of holotype in life

Dorsal surface of head dark brown. Transverse bands on dorsal surface of head indistinct. Lateral surfaces of head brownish grey. A distinct black stripe from posteroinferior eye to tympanum region on each side. Upper lips brownish grey, lower lips white. Oral cavity, inner lips, and tongue pale pink.

Dorsal surface of body dark brown. A creamy yellow moderately jagged dorsolateral stripe on each side of body from occipital region to pelvis. Some indistinct black patterns between two dorsolateral stripes. Some creamy yellow spots scattered below dorsolateral stripe on each side of body. Dorsal surfaces of limbs brown with dark transverse bands. Dorsal surface of tail brownish grey with indistinct dark transverse bands.

Ventral surface of head white. A pale yellow gular spot on posterior central part of ventral head, some short black stripes mostly on anterior and sides of gular spot. Ventral surfaces of body, limbs, and tail white with no pattern.

Variations

The variations of metrical characteristics of the type series are provided in Table 3. Other variations are as follows: the dorsal colour are paler, the patterns between two dorsolateral stripes are more distinct, the transverse bands on the dorsal surface of head are more distinct, and the dorsolateral stripes are indistinct, pale grey or yellow anteriorly and pale grey posteriorly in the female paratypes; the gular spot present in one female paratype and absent in another female paratype; in addition, the short black stripes on ventral surface of head are more indistinct in all paratypes.

Table 3.
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Morphological data of the type series of Diploderma donglangense sp. nov. Morphometric measurements are in the unit of mm. For measurement and count methods and abbreviations, see the Materials and methods.

KIZ2022057 holotype ♂ KIZ2022058 paratype ♀ KIZ2022059 paratype ♀ KIZ2022060 paratype ♂ KIZ2022061 paratype ♀
SVL 52.8 51.2 56.4 44.9 47.2
TAL 93.1 81.4 90.5 84.0 89.1
HL 16.9 16.1 17.3 14.6 15.0
HW 11.7 11.8 12.2 10.4 10.7
HD 9.3 9.6 9.7 8.4 8.5
SEL 5.6 5.5 6.1 5.3 5.3
FLL 24.4 24.1 24.6 21.2 23.8
HLL 38.8 40.1 39.2 33.6 35.4
T4L 9.0 9.2 9.4 8.0 9.5
TRL 23.5 23.0 25.9 19.8 20.8
TAL/SVL 1.76 1.59 1.60 1.87 1.89
SEL/HL 0.33 0.34 0.35 0.36 0.35
HW/HL 0.69 0.73 0.71 0.71 0.71
HD/HW 0.79 0.81 0.80 0.81 0.79
FLL/SVL 0.46 0.47 0.44 0.47 0.50
HLL/SVL 0.73 0.78 0.70 0.75 0.75
TRL/SVL 0.45 0.45 0.46 0.44 0.44
SL 10/9 9/9 8/9 8/8 10/10
IL 11/11 9/10 10/10 10/9 10/9
NSL 2/2 2/2 2/1 1/1 2/1
MD 40 37 43 39 41
F4S 16/16 13/14 17/17 14/15 16/15
T4S 24/23 19/20 21/21 22/22 24/21
SOR 5/4 5/5 4/4 5/5 5/5
VN 63 56 60 61 55

Comparisons

Diploderma donglangense sp. nov. differs from D. brevipes, D. chapaense, D. fasciatum, D. hamptoni, D. luei, D. makii, D. menghaiense, D. micangshanense, D. ngoclinense, D. polygonatum, D. swinhonis, and D. yunnanense by the presence of a transverse gular fold (vs. absence).

Diploderma donglangense sp. nov. differs from D. dymondi, D. panlong, D. slowinskii, D. varcoae, and D. swild by having concealed tympana (vs. exposed).

Diploderma donglangense sp. nov. differs from D. drukdaypo, D. flaviceps, D. shuoquense, D. splendidum, and D. vela by the presence of a distinct gular spot in males in life (vs. absence).

Diploderma donglangense sp. nov. differs from D. aorun, D. batangense, D. bowoense, D. brevicauda, D. daochengense, D. flavilabre, D. formosgulae, D. iadinum, D. laeviventre, D. limingensis, D. xinlongense, D. yangi, D. yongshengense, D. yulongense, and D. zhaoermii by having a pale yellow gular spot in males in life (vs. chartreuse, blue, green, lilac, orange, or yellowish white).

Diploderma donglangense sp. nov. differs from D. angustelinea by the presence of short black stripes on ventral head (vs. absence); from D. grahami by having relatively longer hind limbs (HLL/SVL 0.70–0.78 vs. 0.61), having a distinct transverse gular fold (vs. feeble), and the presence of dorsolateral stripes (vs. absence); from D. kangdingense by the absence of skin folds under nuchal and dorsal crests in males (vs. presence) and having white ventrolateral surface of body in males in life (vs. yellow); from D. panchi by having a relatively longer tail in females (TAL/SVL 1.59–1.89 vs. 1.42–1.52), having relatively longer hind limbs in females (HLL/SVL 0.70–0.78 vs. 0.60–0.66), and the presence of short black stripes on ventral head (vs. absence); and from D. qilin by having short black stripes on ventral head (vs. vermiculate stripes) and having moderately jagged dorsolateral stripes in males (vs. strongly jagged).

Diploderma donglangense sp. nov. differs from Diploderma danbaense sp. nov. by the presence of a distinct gular spot in males in life (vs. absence), the absence of reticulate pattern on ventral head (vs. presence), having moderately jagged dorsolateral stripes in males (vs. strongly jagged), the absence of distinct, dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum (vs. absence), and the absence of skin folds under nuchal and dorsal crests in males (vs. presence).

Figure 7. 

Dorsal view (upper) and ventral view (lower) of the type series of Diploderma donglangense sp. nov. in preservative.

Figure 8. 

Holotype (KIZ2022057) of Diploderma donglangense sp. nov. in life A dorsal view B lateral view C ventral view D close-up view of the dorsolateral side of the head E close-up view of the ventral side of the head F close-up view of the oral cavity.

Figure 9. 

Paratypes of Diploderma donglangense sp. nov. in life A, B the female paratype KIZ2022058 C, D the female paratype KIZ2022059 E, F dorsolateral the male paratype KIZ2022060.

Distribution

This species is currently known only from its type locality in Donglang Township, Muli County, Liangshan Prefecture, Sichuan Province, China (Fig. 1).

Natural history

This species is terrestrial, inhabiting the hot-dry valley of the upper Shuiluo River, which is a tributary of the Jinsha River. There are many thorny shrubs and some rock piles at the type locality (Fig. 16C, D). All specimens were collected between 1 and 3 p.m. when they were basking on rock piles.

Diploderma jiulongense sp. nov.

https://zoobank.org/A60F9BDB-0363-41A2-A9B9-CE7F6E964B56
Figs 10, 11, 12

Type material

Holotype. KIZ2022086, adult male, collected on 8 August 2022 by Mian Hou from Yandai Town, Jiulong County, Ganzi Prefecture, Sichuan Province, China (28°28'39"N, 101°44'57"E, 1680 m elevation).

Paratypes. KIZ2022087 and KIZ2022101, two adult females; KIZ2022099–KIZ2022100, two adult males, collecting information all the same as the holotype.

Etymology

The specific epithet refers to Jiulong County, where the new species was discovered.

Diagnosis

Diploderma jiulongense sp. nov. can be diagnosed from congeners by a combination of the following morphological characteristics: (1) body size moderate, SVL 44.5–51.4 mm in adult males, 61.7–64.4 in adult females; (2) tail very long, TAL/SVL 2.57–2.71 in adult males, 2.33–2.44 in adult females; (3) limbs very long, FLL/SVL 0.50–0.53 in adult males, 0.46–0.47 in adult females, HLL/SVL 0.86–0.94 in adult males, 0.79–0.82 in adult females; (4) head moderately long, HW/HL 0.68–0.73 in adult males, 0.71–0.72 in adult females; (5) MD 36–40; (6) F4S 16–19, T4S 23–27; (7) tympanum concealed; (8) nuchal and dorsal crests continuous, feebly developed, no skin fold under nuchal and dorsal crests; (9) distinct transverse gular fold present; (10) ventral scales of head heterogeneous in size, ones on centre of gular pouch largest, all strongly keeled; (11) ventral scales of body strongly keeled; (12) gular spot present in males, indistinct or absent in females, pale yellow in life; (13) dorsolateral stripes distinct in males, narrow and smooth edged, bright yellow in life; (14) no radial stripes around eye; (15) oral cavity and inner lips pinkish white, tongue pale flesh colour in life.

Description of holotype

Adult male, SVL 51.4 mm; tail very long, TAL 137.1 mm, TAL/SVL 2.67; limbs very long, FLL 27.0 mm on left side, FLL/SVL 0.53, HLL 48.1 mm on left, HLL/SVL 0.94. Head moderately long, HW/HL 0.68, HD/HW 0.86; snout moderately long, SEL/HL 0.37. Rostral elongated, bordered by six small postrostral scales; dorsal head scales heterogeneous, all strongly keeled; distinct Y-shaped ridge on dorsal snout. Nasal oval, separated from first supralabial by single row of scales on each side; loreals small, keeled; suborbital scale rows 4/4, keeled; canthus rostralis elongated, greatly overlapping with each other; enlarged, keeled scales forming single lateral ridge from posteroinferior eye to posterosuperior tympanum on each side; tympanum concealed under scales; SL 8/8, smooth. Mental pentagonal; IL 9/9; enlarged chin shields 4/5, smooth, first one contacting IL on each side, remaining ones separated from IL by two rows of small scales on each side; ventral head scales heterogeneous in size, ones on centre of gular pouch largest, all strongly keeled; distinct transverse gular fold present; gular pouch weakly developed.

Distinct shoulder fold present; dorsal body scales heterogeneous in size and shape, all keeled, tip pointing backwards; axillary scales much smaller than remaining dorsals; enlarged dorsal scales roughly forming four longitudinal rows on each side of body. Nuchal and dorsal crests feebly developed, continuous; no skin fold under nuchal and dorsal crest; MD 36. Dorsal limb scales strongly keeled, mostly homogeneous, except a few enlarged, conical scales on postaxial thighs; F4S 18/19, T4S 27/27. Ventral body scales approximately parallel, homogeneous, all strongly keeled, VN 54. Ventral limb scales parallel, small on upper arms and thighs and larger on forearms and crus, all strongly keeled. Tail scales all strongly keeled, ventral tail scales slightly larger than dorsal tail scales.

Colouration of holotype in life

Dorsal surface of head dark grey with no transverse bands. Lateral surfaces of head grey, a greyish white suborbital stripe extending from nasal scale to rictus on each side. Upper lips brownish grey, lower lips greyish white. No radial stripes around eyes. Oral cavity and inner lips pinkish white, tongue pale flesh colour.

Dorsal surface of body brownish black. A narrow, bright yellow, smooth-edged, dorsolateral stripe on each side of body from occipital region to pelvis. Some black inverted triangular patterns between the two dorsolateral stripes. Some bright yellow spots scattered below dorsolateral stripe on each side of body. Dorsal surfaces of limbs dark grey. Dark transverse bands on dorsal surfaces of limbs very indistinct. Dorsal surface of tail brownish grey with some indistinct dark transverse bands.

Ventral surface of head greyish white. A pale yellow gular spot present on posterior central part of ventral head, no stripes on ventral head. Ventral surfaces of body, limbs, and tail greyish white with no pattern.

Variations

The variations of metrical characteristics of the type series are provided in Table 4. Other variations are as follows: the dorsal colour is brownish red or brick red, the transverse bands on the limbs are more distinct, the dorsolateral stripes are indistinct, pale grey or yellow anteriorly and pale grey posteriorly, and the gular spot indistinct or absent in the female paratypes.

Table 4.
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Morphological data of the type series of Diploderma jiulongense sp. nov. Morphometric measurements are in the unit of mm. For measurement and count methods and abbreviations, see the Materials and methods.

KIZ2022086 holotype ♂ KIZ2022087 paratype ♀ KIZ2022099 paratype ♂ KIZ2022100 paratype ♂ KIZ2022101 paratype ♀
SVL 51.4 61.7 47.8 44.5 64.4
TAL 137.1 143.5 129.7 114.3 157.4
HL 17.4 18.0 14.5 13.4 18.5
HW 11.9 13.0 10.6 9.5 13.1
HD 10.2 10.5 8.6 7.9 10.3
SEL 6.4 6.5 5.6 4.6 7.0
FLL 27.0 28.7 24.9 22.3 29.8
HLL 48.1 50.6 40.9 38.2 51.1
T4L 12.8 12.9 10.1 10.0 12.6
TRL 21.9 31.0 21.6 20.6 31.2
TAL/SVL 2.67 2.33 2.71 2.57 2.44
SEL/HL 0.37 0.36 0.39 0.34 0.38
HW/HL 0.68 0.72 0.73 0.71 0.71
HD/HW 0.86 0.86 0.81 0.83 0.79
FLL/SVL 0.53 0.47 0.52 0.50 0.46
HLL/SVL 0.94 0.82 0.86 0.86 0.79
TRL/SVL 0.43 0.50 0.45 0.46 0.48
SL 8/8 8/8 8/8 9/9 8/9
IL 9/9 9/9 10/10 9/8 10/9
NSL 1/1 1/1 1/1 1/1 2/2
MD 36 40 36 37 36
F4S 18/19 17/16 16/16 16/18 17/17
T4S 27/27 25/25 23/23 23/- 24/23
SOR 4/4 3/4 4/4 3/4 3/3
VN 54 52 54 56 55

Comparisons

Diploderma jiulongense sp. nov. differs from D. brevipes, D. chapaense, D. fasciatum, D. hamptoni, D. luei, D. makii, D. menghaiense, D. micangshanense, D. ngoclinense, D. polygonatum, D. swinhonis, and D. yunnanense by the presence of a transverse gular fold (vs. absence).

Diploderma jiulongense sp. nov. differs from D. dymondi, D. panlong, D. slowinskii, D. varcoae, and D. swild by having concealed tympana (vs. exposed).

Diploderma jiulongense sp. nov. differs from D. drukdaypo, D. flaviceps, D. shuoquense, D. splendidum, and D. vela by the presence of a distinct gular spot in males in life (vs. absence).

Diploderma jiulongense sp. nov. differs from D. aorun, D. batangense, D. bowoense, D. brevicauda, D. daochengense, D. flavilabre, D. formosgulae, D. iadinum, D. laeviventre, D. limingensis, D. xinlongense, D. yangi, D. yongshengense, D. yulongense, and D. zhaoermii by having a pale yellow gular spot in males in life (vs. chartreuse, blue, green, lilac, orange, or yellowish white).

Diploderma jiulongense sp. nov. differs from D. grahami by having a relatively much longer tail (TAL/SVL 2.33–2.71 vs. 1.64), having relatively longer hind limbs (HLL/SVL 0.82–0.94 vs. 0.61), having a distinct transverse gular fold (vs. feeble), and the presence of dorsolateral stripes (vs. absence); from D. kangdingense by having a relatively longer tail (TAL/SVL 2.33–2.71 vs. 1.56–2.27), the absence of skin folds under nuchal and dorsal crests in males (vs. presence), and having greyish white ventrolateral surface of body in males in life (vs. yellow); from D. panchi by having a relatively longer tail in females (TAL/SVL 2.33–2.44 vs. 1.42–1.52), having relatively longer hind limbs in females (HLL/SVL 0.79–0.82 vs. 0.60–0.66), and the presence of short black stripes on ventral head (vs. absence); and from D. qilin by having a relatively longer tail (TAL/SVL 2.33–2.71 vs. 1.74–2.18), having short black stripes on ventral head (vs. vermiculate stripes) and having smooth edged dorsolateral stripes in males (vs. strongly jagged).

Diploderma jiulongense sp. nov. is phylogenetically sister to and most similar in morphology characteristic and colouration to D. angustelinea; however, Diploderma jiulongense sp. nov. can be differentiated from the latter by having a relatively longer tail (TAL/SVL 2.57–2.71 vs. 2.30–2.49 in males, 2.33–2.44 vs. 1.94–2.22 in females), having relatively longer fore-limbs (FLL/SVL 0.50–0.53 vs. 0.46–0.47 in males, 0.46–0.47 vs. 0.41–0.46 in females), having relatively longer hind limbs (HLL/SVL 0.86–0.94 vs. 0.71–0.80 in males, 0.79–0.82 vs. 0.74–0.79 in females), having a greater ratio of head depth to head width (HD/HW 0.81–0.86 vs. 0.73–0.79 in males, 0.79–0.86 vs. 0.73–0.79 in females), and having smooth edged dorsolateral stripes in males (vs. weakly jagged).

Diploderma jiulongense sp. nov. differs from Diploderma danbaense sp. nov. by having a relatively longer tail (TAL/SVL 2.33–2.71 vs. 1.55–1.78), the presence of a distinct gular spot in males in life (vs. absence), the absence of reticulate pattern on ventral head (vs. presence), having smooth edged dorsolateral stripes in males (vs. strongly jagged), the absence of distinct, dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum (vs. absence), and the absence of skin folds under nuchal and dorsal crests in males (vs. presence).

Diploderma jiulongense sp. nov. differs from Diploderma donglangense sp. nov. by having a relatively longer tail (TAL/SVL 2.33–2.71 vs. 1.59–1.89), the absence of short black stripes on ventral head (vs. presence), and having smooth edged dorsolateral stripes in males (vs. moderately jagged).

Figure 10. 

Dorsal view (upper) and ventral view (lower) of the type series of Diploderma jiulongense sp. nov. in preservative.

Figure 11. 

Holotype (KIZ2022086) of Diploderma jiulongense sp. nov. in life A dorsal view B lateral view C ventral view D close-up view of the dorsolateral side of the head E close-up view of the ventral side of the head F close-up view of the oral cavity.

Figure 12. 

Paratypes of Diploderma jiulongense sp. nov. in life A, B the female paratype KIZ2022087 C, D the male paratype KIZ2022099 E, F the female paratype KIZ2022101.

Distribution

This species is currently known only from its type locality in Yandai Town, Jiulong County, Ganzi Prefecture, Sichuan Province, China (Fig. 1).

Natural history

All specimens were collected between 9 and 12 a.m. in bushes or grass in the Yalong River valley. There are a few trees and many rocks at the type locality (Fig. 16E, F).

Diploderma tachengense sp. nov.

https://zoobank.org/9FAC8DF8-7A21-48C7-B6E3-3463B16347B6
Figs 13, 14, 15

Type material

Holotype. KIZ2022028, adult male, collected on 6 July 2022 by Shuo Liu from Kena Village, Tacheng Town, Weixi County, Diqing Prefecture, Yunnan Province, China (27°34'12"N, 99°20'36"E, 2180 m elevation).

Paratypes. KIZ2022027, adult male; KIZ2022029, adult female; KIZ2022038, subadult male; and KIZ2022039, subadult female; collecting information all the same as the holotype.

Etymology

The specific epithet refers to Tacheng Town, where the new species was discovered.

Diagnosis

Diploderma tachengense sp. nov. can be diagnosed from congeners by a combination of the following morphological characteristics: (1) body size moderate, SVL 55.2–55.7 mm in adult males, 65.4 mm in adult female; (2) tail moderately long, TAL/SVL 1.88–1.89 in adult males, 1.56 in adult female; (3) limbs moderately long, FLL/SVL 0.45–0.47 in adult males, 0.46 in adult female, HLL/SVL 0.76–0.78 in adult males, 0.70 in adult female; (4) head relatively robust, HW/HL 0.73–0.80 in adult males, 0.74 in adult female; (5) MD 38–44; (6) F4S 15–19, T4S 20–24; (7) tympanum concealed; (8) nuchal and dorsal crests continuous, feebly developed, no skin fold under nuchal and dorsal crests; (9) distinct transverse gular fold present; (10) ventral scales of head homogeneous, feebly keeled; (11) ventral scales of body strongly keeled; (12) gular spot present in both sexes, pale yellow in life; (13) dorsolateral stripes distinct in males, strongly jagged, pale yellow in life; (14) radial stripes around eye relatively distinct; (15) oral cavity and inner lips pinkish white, tongue pink in life.

Description of holotype

Adult male, SVL 55.2 mm; tail moderately long, TAL 104.5 mm, TAL/SVL 1.89; limbs moderately long, FLL 25.9 mm on left side, FLL/SVL 0.47, HLL 41.7 mm on left, HLL/SVL 0.76. Head relatively robust, HW/HL 0.73, HD/HW 0.87; snout moderately long, SEL/HL 0.36. Rostral elongated, bordered by six small postrostral scales; dorsal head scales heterogeneous, all strongly keeled; distinct Y-shaped ridge on dorsal snout. Nasal oval, separated from first supralabial by single row of scales on each side; loreals small, keeled; suborbital scale rows 4/4, keeled; canthus rostralis elongated, greatly overlapping with each other; enlarged, keeled scales forming distinct single lateral ridge from posteroinferior eye to posterosuperior tympanum on each side; tympanum concealed under scales; SL 9/9, feebly keeled. Mental approximately triangular; IL 10/10; enlarged chin shields 4/4, smooth, first one contacting IL on each side, second ones separated from IL by one row of small scales on each side, remaining ones separated from IL by two rows of small scales on each side; ventral head scales homogeneous in size, feebly keeled; distinct transverse gular fold present; gular pouch weakly developed.

Distinct shoulder fold present; dorsal body scales heterogeneous in size and shape, all keeled, tip pointing backwards; axillary scales much smaller than remaining dorsals; enlarged dorsal scales forming one intermittent longitudinal row between dorsal crest and dorsolateral stripe on each side, remaining enlarged dorsal scales irregularly scattered on each side of body. Nuchal and dorsal crests feebly developed, continuous; no skin fold under nuchal and dorsal crests; MD 43. Dorsal limb scales strongly keeled, mostly homogeneous, except a few enlarged, conical scales on postaxial thighs; F4S 16/17, T4S 21/22. Ventral body scales approximately parallel, homogeneous, all strongly keeled, VN 55. Ventral limb scales parallel, small on upper arms and thighs and larger on forearms and crus, all strongly keeled. Tail scales all strongly keeled, ventral tail scales slightly larger than dorsal tail scales.

Colouration of holotype in life

Dorsal surface of head dark brown with indistinct transverse bands. Lateral surfaces of head brownish yellow. Upper lips brownish grey, lower lips white. Distinct radial stripes around eye on each side. Oral cavity and inner lips pinkish white, tongue pink.

Dorsal surface of body brownish black. A pale yellow strongly jagged dorsolateral stripe on each side of body from occipital region to pelvis. Several pale yellow spots scattered below dorsolateral stripe on each side of body. Dorsal surfaces of limbs brown. Distinct dark transverse bands present on dorsal surfaces of limbs. Dorsal surface of tail brown with indistinct dark transverse bands.

Ventral surface of head white. A pale yellow gular spot present on posterior central part of ventral head, indistinct dark stripes on other parts of ventral head. Ventral surface of body white with no pattern, ventral surfaces of limbs and tail reddish white.

Figure 13. 

Dorsal view (upper) and ventral view (lower) of the type series of Diploderma tachengense sp. nov. in preservative.

Figure 14. 

Holotype (KIZ2022028) of Diploderma tachengense sp. nov. in life A dorsal view B lateral view C ventral view D close-up view of the dorsolateral side of the head E close-up view of the ventral side of the head F close-up view of the oral cavity.

Figure 15. 

Paratypes of Diploderma tachengense sp. nov. in life A, B the female paratype KIZ2022029 C, D the subadult male paratype KIZ2022038 E, F the subadult female paratype KIZ2022039.

Variations

The variations of metrical characteristics of the type series are provided in Table 5. Other variations are as follows: the dorsal colour is paler, there are some dark inverted triangular patterns between the two dorsolateral stripes, the radial stripes around eyes are more indistinct, and the dark stripes on ventral head are more distinct in all paratypes; in addition, the dorsolateral stripes are indistinct, yellowish white in the female paratypes.

Table 5.
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Morphological data of the type series of Diploderma tachengense sp. nov. Morphometric measurements are in the unit of mm. For measurement and count methods and abbreviations, see the Materials and methods.

KIZ2022028 holotype ♂ KIZ2022027 paratype ♂ KIZ2022029 paratype ♀ KIZ2022038 paratype subadult ♂ KIZ2022039 paratype subadult ♀
SVL 55.2 55.7 65.4 42.7 40.6
TAL 104.5 104.8 102.1 81.6 73.6
HL 17.5 17.7 18.4 13.9 12.6
HW 12.7 14.1 13.6 10.3 9.2
HD 11.1 10.9 11.8 8.8 7.7
SEL 6.3 6.2 6.4 5.1 4.5
FLL 25.9 25.0 30.2 21.8 19.6
HLL 41.7 43.6 45.6 36.4 33.1
T4L 10.2 9.4 10.9 9.0 8.5
TRL 24.1 25.1 33.1 18.7 17.9
TAL/SVL 1.89 1.88 1.56 1.91 1.81
SEL/HL 0.36 0.35 0.35 0.37 0.36
HW/HL 0.73 0.80 0.74 0.74 0.73
HD/HW 0.87 0.77 0.87 0.85 0.84
FLL/SVL 0.47 0.45 0.46 0.51 0.48
HLL/SVL 0.76 0.78 0.70 0.85 0.82
TRL/SVL 0.44 0.45 0.51 0.44 0.44
SL 9/9 8/8 9/9 9/9 8/9
IL 10/10 9/9 11/10 9/10 10/9
NSL 1/1 1/1 1/1 1/1 1/1
MD 43 38 39 44 38
F4S 16/17 16/15 19/19 16/16 17/16
T4S 21/22 21/21 24/23 20/20 21/20
SOR 4/4 4/4 4/4 4/4 4/4
VN 55 51 56 51 53

Comparisons

Diploderma tachengense sp. nov. differs from D. brevipes, D. chapaense, D. fasciatum, D. hamptoni, D. luei, D. makii, D. menghaiense, D. micangshanense, D. ngoclinense, D. polygonatum, D. swinhonis, and D. yunnanense by the presence of a transverse gular fold (vs. absence).

Diploderma tachengense sp. nov. differs from D. dymondi, D. panlong, D. slowinskii, D. varcoae, and D. swild by having concealed tympana (vs. exposed).

Diploderma tachengense sp. nov. differs from D. drukdaypo, D. flaviceps, D. shuoquense, D. splendidum, and D. vela by the presence of a distinct gular spot in males in life (vs. absence).

Diploderma tachengense sp. nov. differs from D. batangense, D. bowoense, D. brevicauda, D. daochengense, D. flavilabre, D. formosgulae, D. iadinum, D. laeviventre, D. limingensis, D. xinlongense, D. yangi, D. yongshengense, D. yulongense, and D. zhaoermii by having a pale yellow gular spot in males in life (vs. chartreuse, blue, green, lilac, orange, or yellowish white).

Diploderma tachengense sp. nov. differs from D. angustelinea by wide strongly jagged dorsolateral stripes in males (vs. narrow, feebly jagged); D. grahami by having relatively longer hind limbs (HLL/SVL 0.70–0.78 vs. 0.61), having a distinct transverse gular fold (vs. feeble), and the presence of dorsolateral stripes (vs. absence); from D. kangdingense by the absence of skin folds under nuchal and dorsal crests in males (vs. presence), and having white ventrolateral surface of body in males in life (vs. yellow); from D. panchi by having relatively longer hind limbs in females (HLL/SVL 0.70 vs. 0.60–0.66), the presence of a distinct gular spot in females in life (vs. absence), and the presence of black stripes on ventral head (vs. absence); and from D. qilin by having a relatively shorter tail (TAL/SVL 1.88–1.89 vs. 2.01–2.18 in males, 1.56 vs. 1.74–2.00 in females) and having a pink tongue (vs. pale flesh coloured).

Diploderma tachengense sp. nov. is phylogenetically sister to D. aorun; however, Diploderma tachengense sp. nov. can be differentiated from the latter by having a pale yellow gular spot in both sexes in life (vs. blue in both sexes), having a relatively much shorter tail (TAL/SVL 1.88–1.89 vs. 2.12–2.21 in males, 1.56 vs. 1.91–2.08 in females), having a greater ratio of head width to head length (HW/HL 0.73–0.80 vs. 0.68–0.73 in males, 0.74 vs. 0.67–0.70 in females), and the absence of skin folds under nuchal and dorsal crests in males (vs. presence).

Diploderma tachengense sp. nov. differs from Diploderma danbaense sp. nov. by the presence of a distinct gular spot in both sexes in life (vs. absence), the presence of distinct radial stripes around eyes (vs. absence), the absence of distinct, dark, hollow, approximately rhomboid patterns between dorsolateral stripes on dorsum (vs. absence), and the absence of skin folds under nuchal and dorsal crests in males (vs. presence).

Diploderma tachengense sp. nov. differs from Diploderma donglangense sp. nov. by having vermiculate stripes on ventral head (vs. short black stripes), having strongly jagged dorsolateral stripes in males (vs. moderately jagged), and the presence of distinct radial stripes around eyes (vs. absence).

Diploderma tachengense sp. nov. differs from Diploderma jiulongense sp. nov. by having a relatively shorter tail (TAL/SVL 1.56–1.89 vs. 2.33–2.71), the presence of vermiculate stripes on ventral head (vs. absence), and having wide, strongly jagged dorsolateral stripes in males (vs. narrow, smooth edged).

Distribution

This species is currently known only from its type locality in Tacheng Town, Weixi County, Diqing Prefecture, Yunnan Province, China (Fig. 1).

Natural history

All specimens were collected between 1 and 5 p.m. on the ground in the secondary coniferous forest (Fig. 16G, H) beside the Lapu River, which is a tributary of the Jinsha River.

Figure 16. 

Habitats of the new species A distant view of the type locality of Diploderma danbaense sp. nov. B close view of the type locality of Diploderma danbaense sp. nov. C distant view of the type locality of Diploderma donglangense sp. nov. D close view of the type locality of Diploderma donglangense sp. nov. E distant view of the type locality of Diploderma jiulongense sp. nov. F close view of the type locality of Diploderma jiulongense sp. nov G distant view of the type locality of Diploderma tachengense sp. nov. H close view of the type locality of Diploderma tachengense sp. nov.

Discussion

With a rapidly increasing number of species, the genus Diploderma consists of 42 species in total, whereof nearly half were described in the last five years, and more than one third were described in the last three years (e.g., Cai et al. 2022; Liu et al. 2022; Uetz et al. 2022). Together with the four new species described in this study, we bring the total number of species in this genus to 46.

Among the four new species described in this study, Diploderma jiulongense sp. nov. and Diploderma tachengense sp. nov. have relatively small genetic distances (2.9% and 2.8%) from the closely related species D. angustelinea and D. aorun. However, there are significant morphological differences between Diploderma jiulongense sp. nov. and D. angustelinea and between Diploderma tachengense sp. nov. and D. aorun. Diploderma jiulongense sp. nov. differs from D. angustelinea by having a relatively much longer tail, relatively longer fore-limbs, relatively longer hind limbs, and having smooth edged dorsolateral stripes in males. Diploderma tachengense sp. nov. differs from D. aorun by having an obviously differently coloured gular spot in life, having a relatively shorter tail, and the absence of skin folds under nuchal and dorsal crests in males in life. In addition, the genetic distances between them and their closely related species are both greater than that (2.6%) between the two recognized species D. drukdaypo and D. vela. Although the genetic distance is small between D. drukdaypo and D. vela, they have distinct and stable morphological differences (i.e., relative limb length, relative tail length, and keeling of ventral body scales; Wang et al. 2015, 2019b). Thus, they are regarded as two distinct species each that occupy distinct sections of the Mekong River and live at different elevations (Wang et al. 2021b). Therefore, taking the case of the species pair of D. drukdaypo and D. vela as the calibration, Diploderma jiulongense sp. nov. and Diploderma tachengense sp. nov. also have surpassed intraspecific variation level and reached a specific status.

Another new species described in this study, Diploderma danbaense sp. nov., is at first glance similar in morphology to D. flaviceps. To compare morphological characteristics between Diploderma danbaense sp. nov. and D. flaviceps, we examined 27 topotypic or near-topotypic specimens of D. flaviceps (Suppl. material 2), and found several non-overlapping morphological characteristics (i.e., relative tail length, relative hind limb length, and relative head width) between both taxa (see Table 6). In addition, the genetic distance (4.4%) between Diploderma danbaense sp. nov. and D. flaviceps is approximate to that (4.1%) between D. brevicauda and D. qilin, greater than that (3.2%) between D. formosgulae and D. drukdaypo and that (3.3%) between D. formosgulae and D. vela, and much greater than that (2.6%) between D. drukdaypo and D. vela. Diploderma brevicauda and D. qilin are similar in morphology, according to Wang et al. (2021b), there are only differences in relative tail length, relative hind limb length and oral cavity colour between them, and these differences are considered sufficient to distinguish different species (Wang et al. 2021b).

Table 6.
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Comparison between Diploderma danbaense sp. nov. and D. flaviceps. For morphological abbreviations see the Materials and methods. Detailed measurements and pholidosis data for each examined specimen of D. flaviceps are given in Suppl. material 2.

Diploderma danbaense sp. nov. Diploderma flaviceps
♂ (n = 4) ♀ (n = 1) ♂ (n = 18) ♀ (n = 9)
SVL 68.7–77.0 76.6 60.5–75.6 53.4–67.1
TAL 112.7–130.0 119.1 117.7–151.0 103.7–127.0
HL 21.4–26.6 23.5 18.9–24.9 16.3–20.5
HW 15.0–17.9 14.7 14.6–20.6 12.6–15.6
HD 12.8–15.0 12.8 10.7–14.8 9.6–12.5
SEL 7.5–9.9 8.4 5.9–8.9 5.2–6.8
FLL 28.7–33.1 33.6 28.8–36.1 24.3–31.6
HLL 49.1–55.2 52.1 48.1–60.0 39.2–50.9
T4L 12.2–12.8 12.5 11.4–14.9 9.6–11.7
TRL 29.2–33.2 34.5 26.7–34.6 24.3–33.0
TAL/SVL 1.61–1.78 1.55 1.88–2.09 1.73–2.17
HL/SVL 0.31–0.35 0.31 0.31–0.34 0.29–0.33
HW/HL 0.66–0.75 0.63 0.76–0.84 0.71–0.78
HD/HW 0.80–0.85 0.87 0.70–0.78 0.75–0.83
FLL/SVL 0.41–0.47 0.44 0.44–0.49 0.43–0.49
HLL/SVL 0.66–0.70 0.65 0.72–0.80 0.70–0.81
T4L/SVL 0.16–0.18 0.16 0.18–0.21 0.17–0.21
SL 9–10 10–11 8–11 9–11
IL 10–11 11 10–13 10–12
NSL 1–2 2 1–2 1–2
MD 47–53 58 40–55 46–54
F4S 16–20 19–20 16–21 16–19
T4S 21–25 24–26 22–26 22–27
SOR 4–5 4–5 3–5 3–5

Allopatric distribution across the Hengduan Mountain Region have been suggested to create reproductive isolation through geographic isolation (Wang et al. 2015, 2021a; Dong et al. 2020). Wang et al. (2021b) considered that D. drukdaypo may represent a recently diverged species from the D. vela lineage, adapted to higher-elevated habitats in the upper Mekong River valley, and suggested that further studies are needed to gain a better understanding of gene flow and speciation mechanisms between these two sister species. Although D. formosgulae is morphologically similar and genetically close to D. drukdaypo and D. vela, Wang et al. (2021a) stated that D. formosgulae possesses an allopatric distribution with the latter two species and represents a recently diverged lineage, evolutionarily distinct. Likewise, Diploderma danbaense sp. nov. also possesses an allopatric distribution with D. flaviceps and may represent a recently diverged species, adapted to higher-elevated habitats in the upper Dadu River valley.

To further confirm the validity of these three new species, we conducted detailed surveys in the field to verify whether there is geographical isolation between these three new species and their closely related relatives. At first, the type locality of Diploderma tachengense sp. nov. (at 2180 m elevation) is approximately 100 km downstream along the Jinsha River from the type locality of D. aorun (at 2198 m elevation). Although the altitudinal ranges are similar, their habitats are completely different, one is a hot-dry valley of the Jinsha River, the other is a distant forested region. Therefore, there is a clear geographical and ecological isolation between the populations of Diploderma tachengense sp. nov. and D. aorun. Further, the type locality of Diploderma jiulongense sp. nov. (at 1680 m elevation) is approximately 130 km downstream along the Yalong River from the type locality of D. angustelinea (at 2017 m elevation). Although the linear distance between the two localities is short, there is a branch of the Daxue Mountain, with the highest peak at approximately 5000 m elevation, between them, serving as a clear geographic barrier. In addition, there is another population of Diploderma (D. cf. daochengense) occupying part valley section of the Yalong River, which bypasses the branch of the Daxue Mountain, between the type localities of Diploderma jiulongense sp. nov. and D. angustelinea. At last, the type locality of Diploderma danbaense sp. nov. (at 2020 m elevation) is approximately 150 km upstream along the Dadu River from the type locality of D. flaviceps (at 1300 m elevation). Although there is no obvious geographical barrier between the two localities except for the altitudinal difference, we have not found any transitional type individuals between Diploderma danbaense sp. nov. and D. flaviceps, nor have we found Diploderma danbaense sp. nov. and D. flaviceps occurring in sympatry. In addition, the relatively great genetic distance (4.4%) between Diploderma danbaense sp. nov. and D. flaviceps also supports an isolation process. A similar situation occurs between the two recognized closely related species D. panlong and D. swild. Through our observation, we found that these two species are similar in morphology and colouration, their habitats and natural history are also similar, as both inhabit forests or forest margins and both are more arboreal than terrestrial, and there is no obvious geographical barrier between them. However, these two species have not been found in sympatry, and there is a great genetic distance between them. We speculate that some historical reasons may have led to the isolation of different populations, thus, they lost genetic exchange and differentiated into different species.

Finally, the mitochondrial ND2 gene is considered to be able to better distinguish different species and has been widely used in phylogenetic analyses of Agamidae (i.e., Zug et al. 2006; Grismer et al. 2016; Ambekar et al. 2020; Liu et al. 2021; Wang et al. 2022a), and the phenomenon of different species with small genetic distances in the ND2 gene is well known from other genera of Agamidae, such as in the genus Sitana Cuvier, where a genetic distance of 3–4% in the ND2 gene is considered sufficient to distinguish different species (Ambekar et al. 2020). This supports our taxonomic actions presented herein, viz. that different populations with relatively small genetic distances in the ND2 gene can represent different species in the Agamidae.

Acknowledgements

Thanks to the curator of Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, for his support of the field survey and taxonomic research. We also thank the editors and reviewers for their comments on the manuscript. This work was supported by Science-Technology Basic Condition Platform from the Ministry of Science and Technology of the People’s Republic of China (2005DKA21402); state theme of Zoological Institute, Russian Academy of Sciences (122031100282-2); the funds from National Science Foundation of China to DQ Rao (NSFC-39570090 and NSFC-31970404); the project of Ministry of Ecology and Environment of China: Investigation and evaluation of amphibian and reptile diversity in Yunling mountains (2019HB2096001006); and National important research and development project: Biodiversity conservation and restoration technology in high mountain and valley regions of southwestern China (2017YFC0505202).

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Supplementary materials

Supplementary material 1 

Uncorrected genetic pairwise distances (p-distances) (%) between species based on the mitochondrial ND2 gene sequences.

Shuo Liu, Mian Hou, Natalia B. Ananjeva, Dingqi Rao

Data type: table (excel document)

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 2 

Detailed morphological measurements and pholidosis characteristics for each examined specimen of Diploderma flaviceps. All measurements are in the unit of mm. “–” indicates not available. For measurement and count methods and abbreviations, see the Materials and methods.

Shuo Liu, Mian Hou, Natalia B. Ananjeva, Dingqi Rao

Data type: table (excel document)

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (42.00 kb)
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