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Diploderma formosgula WANG, GAO, WU, DONG, SHI, QI, SILER & CHE, 2021

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Higher TaxaAgamidae (Draconinae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Vibrant-gulared Mountain Dragon
Chinese: 丽喉龙蜥 (Pinyin: Li Hou Long Xi) 
SynonymDiploderma formosgulae WANG, GAO, WU, DONG, SHI, QI, SILER & CHE in WANG et al. 2021
Japalura flaviceps — ZHAO & YANG 1997: 165
Japalura flaviceps — ZHAO et al. 1999: 111
Japalura flaviceps — YANG & RAO 2008: 200 
DistributionChina (Sichuan: Muli County)

Type locality: Yangla Village, Deqin County, Yunnan Province, China (99.1113˚ E, 28.8905˚ N, elevation 2355m, WGS 84).  
Reproduction 
TypesHolotype: KIZ 044425, adult male, collected by Kai Wang, Zhuoyu Lu, Man Fu, and Xiankun Huang on 17 June 2019.
Paratypes: KIZ 044373, 044417, adult males; KIZ 044418, subadult male; KIZ 044424, 044427, 044428, 044435, juvenile males; KIZ 044375, 044420, 044421, 044423, adult females; KIZ 044429, 044430, 044437, juvenile females. All share the same collection information as the holotype. 
DiagnosisDiagnosis: The new species can be diagnosed based on a combination of the following morphological characters: 1) body size moderate, SVL 55.5–60.9 mm in males, 55.9–60.2 mm in females; 2) tail long, TAL 195.8–223.5% SVL in males, 186.1–194.8% in females; 3) HW 66.2–75.4% HL; 4) HLL 73.0–80.3% SVL; 5) nuchal and dorsal crests developed on moderate skin folds; 6) MD 37–48; 7) F4S 13–16; 8) T4S 18–24; 9) PTS 2–4; 10) PTY 3–6; 11) PRS 4–8; 12) subocular regions and chin pale Pink [Color 242] in most individuals; 13) dorsolateral stripes jagged and Cream Color [Color 12] in males, narrow and Sulphur Yellow [Color 80] or indistinct in females; 14) ventral body uniform Pale Pinkish Buff [Color 3] to Pale Sulfur Yellow [Color 92]; 15) gular spots always present in males, Pinkish Flesh Color [Color 253] to Lilac [Color 222], either as Sulphur Yellow [Color 80] gular spots or Sulphur Yellow [Color 80] reticulated stripes in females (Wang et al. 2021).

Comparisons: The new species has been confused with D. flaviceps, but it can be differentiated by the latter by having a smaller body size in males (SVL 55.5–60.9 mm vs. 68.5–82.1mm) and by the presence of distinct radial stripes around eyes (vs. absence), absence of skin folds under nuchal crests in females (vs. presence), absence of lateral series of dark rhomboid patterns between dorsolateral stripes (vs. presence), and by the presence of colorful gular spots or gular patterns in both sexes (vs. absence).
For species that are phylogenetically closely related but allopatric (D. drukdaypo and D. vela), D. formosgula can be differentiated from the latter two by the presence of colorful gular spots in both sexes (vs. absence in both D. drukdaypo and D. vela). In addition, the new species differ from D. drukdaypo by having distinctively keeled ventral scales of head and body (vs. smooth or feebly keeled) and a much longer tail (TAL 195.8–223.5% SVL in males, 186.1–194.8% in females vs. 153.0–154.4% in males, 132.8–144.0% in females; Fig. 6); and from D. vela by the absence of strongly developed, continuous sail under the vertebral crests in males (vs. presence).
For all remaining recognized species, D. formosgula can be differentiated from all species within the HMR (except D. grahami and D. panchi) by having distinct coloration of gular spots in life (Pinkish Flesh Color [Color 253] to Lilac [Color 222] in males, Sulphur Yellow [Color 80] gular spots or reticulated patterns in females vs. Pale Cyan [Color 157] to Light Caribbean Blue [Color 163] in both sexes of D. aorun; Spectrum Yellow [Color 79] to Dark Spectrum Yellow [Color 78] in D. angustelinea; Light Cyan [Color 157] in both sexes of D. batangense; Light Chrome Orange [Color 76] in males, Dark Spectrum Yellow [Color 78] in females of D. bowoense ; Pale Sulphur Yellow [Color 92] in males, no gular spots in females of D. brevicaudum; Pale Emerald Green [Color 141] to Light Turquoise Green [Color 146] in both sexes of D. flavilabre; Caribbean Blue [Color 168] in males, Medium Greenish Yellow [Color 88] in females of D. iadinum; Light Chrome Orange [Color 76] in both sexes of D. laeviventre; no gular spots in D. panlong, D. slowinskii and D. swild; Light Sulphur Yellow [Color 93] in both sexes of D. qilin; Chartreuse [Color 89] or Opaline Green [Color 106] in both sexes of D. yulongense; and Chartreuse [Color 89] in males, no gular spots in females of D. zhaoermii).
In addition to distinct gular coloration, D. formosgula differs from D. angustelinea by having a shorter tail (TAL 195.8–223.5% SVL in males, 186.1–194.8% in females vs. 230.2–249.1% in males, 194.30–222.3% in females; Fig. 6) and more developed nuchal and dorsal crests in males (vs. feeble); from D. aorun by a distinct coloration of subocular region and chin (pale Pink [Color 242] vs. white); from D. batangense by having a better developed and more conical scales post rictus (PRS 4–9 vs. 0–3); from D. bowoense and D. brevicaudum by having a tendency toward longer tail in males, much longer in females (TAL 195.8–223.5% SVL in males, 186.1–194.8% in females vs. 189.1–201.9% in males, 177.3–178.7% in females for D. bowoense ; 140–184.1% in males, 125.0–159.1% in females of D. brevicaudum; Fig. 6); from D. iadinum by having less robust head (HD 66.9–73.4% HW vs. 74.0–79.9%) and distinct body coloration (Light Buff [Color 2] to Pale Pinkish Buff [Color 3] vs. Yellowish Spectrum Green [Color 128] to Emerald Green [Color 143] in males, Buff [Color 5] to [Pale Greenish Yellow [Color 86] in females); from D. laeviventre by having distinctively keeled ventral scales of head and body (vs. smooth or feebly keeled); from D. slowinskii and D. swild by having concealed tympanum (vs. exposed); from D. yulongense by the absence of green patches on ventrolateral body of males (vs. presence) and by having more conical scales post rictus (PRS 4–9 vs. 1–4); from D. zhaoermii by having a smaller body size (SVL 55.5–60.9 mm in males, 55.9–60.2mm in females vs. 63.24–81.7mm in males, 61.0–75.2mm in females) and less and more scattered dark radial stripes around eyes (vs. more and densely positioned).
For D. grahami, which does not have coloration data in life, the new species differ from the former by having a much longer tail in both sexes (TAL 186.1–223.5% vs. ≤150%), and from the latter by having no granular scales (vs. presence) and by having a distinct transverse gular fold (vs. absence). For D. panchi that has similar gular patterns in females, D. formosgula differs from D. panchi by having a much longer tail (TAL 186.1–194.8% in females vs. 141.8–151.5%).
For the remaining species, D. formosgula differs from D. chapaense, D. fasciatum, D. menghaiense, D. micangshanense, D. varcoae, D. yunnanense, and all species from East Asian islands (D. brevipes, D. luei, D. makii, D. polygonatum, and D. swinhonis) by the presence of a deep transverse gular fold (vs. absence); from D. dymondi by having concealed tympana (vs. exposed); from D. hamptoni by having parallel dorsolateral stripes in males (vs. diagonally oriented) and by having a much deeper transverse gular fold (vs. feeble) (Wang et al. 2021).

Color in life: The background coloration of the dorsal and lateral surfaces of the head is pale Light Flesh Color [Color 250] to pale Pink [Color 242]. Three Sepia [Color 286] to Jet Black [Color 300] transverse ornamentation patterns are present on the dorsal surface of the head: the first one is somewhat irregular shaped between the anterior edge of the orbit, the second one is a cross-band between the midpoint of the orbits, and the third one is a X-shaped pattern between the posterior edge of the orbit. All three of these transverse ornamentation patterns extend laterally and enter the orbit on each side of the head; and together with five additional Jet Black [Color 300] streaks or series of speckles on the lateral surface of the head on each side, they form the radial stripe pattern around the eyes. The color of the upper lips and the superior portion of the corner of the mouth are more saturated. Random Sepia [Color 286] to Jet Black [Color 300] short strakes and speckles are also present on the dorsal and lateral surfaces of the head.
The background coloration of the dorsum is Light Buff [Color 2] to Pale Pinkish Buff [Color 3]. A Sepia [Color 286] to Jet Black [Color 300] reticulated ornamentation pattern is present on the lateral surface of the body, inferior to the dorsolateral stripes. A dorsolateral stripe is present on each side of the vertebral crest on the dorsum, running from the neck to the pelvis. This dorsolateral stripe is strongly jagged, with six rhombus-shaped extensions distributed evenly across the stripe from the shoulder to the pelvis. Such extensions connect dorsally with the counterparts from the other side of the body, forming six transverse streaks across the vertebral crest. The rhombus extensions and their connecting parts of the dorsolateral stripes are Light Buff [Color 2], where the remaining parts of the dorsolateral stripes are muddied with Burnt Umber [Color 48] speckles and patches. The dorsal surfaces of the limbs and tail are Light Buff [Color 2], with evenly distributed Sepia [Color 286] to Jet Black [Color 300] transverse bands or streaks running from the proximal to the distal ends. These streaks get much paler as they approach the distal ends of the fingers, toes, and the tip of the tail.
The ventral surface of the head is white, with Jet Black [Color 300] speckles and short streaks, which is formed by singular rows of black scales. A distinct, triangular shaped gular spot is present at the posterior center of the gular, which is Rose Pink [Color 243] on the peripheral area and Lilac [Color 222] in the center. The ventral surface of the body is uneven Sulphur Yellow [Color 80], which is more saturated towards the anterior portion of each individual scale and paler towards the posterior tip. No other ornamentation is present on the ventral surface of the body. The ventral surfaces of the limbs and the tail are uniform white, except the distal portion, which becomes more brownish (Wang et al. 2021). 
CommentNomenclature: The proper spelling of the species Latin name is Diploderma formosgula, instead of D. formosgulae, which is what the original description used (personal communication with Dr. W. Denzer to K. Wang, pers. comm., 28 Apr 2021 )

Similar species: D. drukdaypo and D. vela

Conservation: Currently, it is known only along a short section of the upper Jinsha River valley, with an estimated range of about 50km in linear distance, and about 350 km2 in area of occupancy. Serious habitat degradation was observed at the type locality, due to both natural causes (i.e. recent flooding from the barrier lake upstream in 2018), as well as anthropogenic causes (i.e. constructions for road repair) (Fig. 11 C1 and C2). In comparison with congeners in the lower reaches of the Jinsha River (i.e. D. aorun, and D. qilin), the population density of D. formosgula is relatively lower. Currently, the new species’ range overlaps with major roads and townships in the area, and the habitat is not protected by any existing nature reserve. Following the IUCN listing criteria B 2b (iii) (area of occupancy estimated less than 500km2, observed continuing decline in quality of habitat), we recommend listing the new species as Endangered (EN), and we recommend listing it as Class II protected for Chinese Wildlife Protecting Act, in the hope this will bring protection to its degrading valley habitats. 
EtymologyThe Latin species epithet formosgula means “beautiful gular”, which describe the vibrant and diagnostic gular spots of the species. 
References
  • Wang, K., Ren, J. L., Jiang, K., Wu, J. W., Yang, C. H., Xu, H. M., ... & Che, J. 2019. Revised distributions of some species in the genus Diploderma (Reptilia: Agamidae) in China. Sichuan J. Zool, 38(5), 481-495
  • Wang, K., Ren, J., Wu, J., Jiang, K., Jin, J., Hou, S., ... & Che, J. 2020. Systematic revision of mountain dragons (Reptilia: Agamidae: Diploderma) in China, with descriptions of six new species and discussion on their conservation. Journal of Zoological Systematics and Evolutionary Research, 59(1), 222-263 (published online 2020)
  • WANG, Kai; , Ke JIANG, Gang PAN, Mian HOU, Cameron D. SILER and Jing CHE 2015. A New Species of Japalura (Squamata: Sauria: Agamidae) from Upper Lancang (Mekong) Valley of Eastern Tibet, China. Asian Herpetological Research 6 (3): 159–168 - get paper here
  • Wang, Kai; Jing Che, Simin Lin, V Deepak, Datta-Roy Aniruddha, Ke Jiang, Jieqiong Jin, Hongman Chen, Cameron D Siler; 2018. Multilocus phylogeny and revised classification for mountain dragons of the genus Japalura s.l. (Reptilia: Agamidae: Draconinae) from Asia. Zoological Journal of the Linnean Society, , zly034, - get paper here
  • WANG, Kai; Ke JIANG, Da-Hu ZOU, Fang YAN, Cameron D. SILER, Jing CHE 2016. Two new species of Japalura (Squamata: Agamidae) from the Hengduan Mountain Range, China. Zoological Research 37(1): 41-56, DOI: 10.13918/j.issn.2095-8137.2016.1.41 - get paper here
  • WANG, KAI; WEI GAO, JIAWEI WU, WENJIE DONG, XIAOGANG FENG, WENJING SHEN, JIEQIONG JIN, XIUDONG SHI, YIN QI, CAMERON D. SILER, JING CHE 2021. Two New Species of Diploderma Hallowell, 1861 (Reptilia: Squamata: Agamidae) from the Hengduan Mountain Region in China and Rediscovery of D. brevicaudum (Manthey, Wolfgang, Hou, Wang, 2012). Zootaxa 4941 (1): 001–032 - get paper here
  • Yang DT, Rao DQ. 2008. Amphibia and Reptilia of Yunnan. Kunming: Yunnan Publishing Group Corporation, 1-411. (in Chinese)
  • Zhao E. M., Zhao K., Zhou K. Y. 1999. Fauna Sinica, Reptilia, Vol. 2, Squamata, Lacertilia. [In Chinese] Beijing: Science Press, 394 pp
  • Zhao, E & Yang, D. 1997. Amphibians and Reptiles of the Hengduan Mountain Region. [in Chinese] Science Press, Beijing, 303 pp.
 
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