Diploderma bowoense, Wang & Gao & Wu & Dong & Feng & Shen & Jin & Shi & Qi & Siler & Che, 2021

Wang, Kai, Gao, Wei, Wu, Jiawei, Dong, Wenjie, Feng, Xiaogang, Shen, Wenjing, Jin, Jieqiong, Shi, Xiudong, Qi, Yin, Siler, Cameron D. & Che, Jing, 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), pp. 1-32 : 14-19

Diploderma bowoense sp. nov. Wang, Gao, Wu, Siler, Che

( Figs. 3 , 6 , 7 , 8 , and 11; Table 4)

LSID urn:lsid:zoobank.org:act:DC266DAA-E8C2-4BB7-84D3-469361331F42

Chresonyms. Japalura flaviceps Zhao & Yang 1997: 165–167 ; Zhao et al. (1999: 111–115), in part; Zhao (2004): 84.

Holotype. KIZ 044700, adult male, from Bowo Village , Muli Tibetan Autonomous County, Sichuan Province, China (101.1569˚ E, 28.8098˚ N, elevation 2328m, WGS 84). Collected by Kai Wang , Zhuoyu Lu , Zhongyang Cireng , and Xiankun Huang on 25 June 2019.

Paratypes. KIZ 044701, adult male; KIZ 044703, 044756, adult females; All share the same collection information as the holotype.

Etymology. The Latin species epithet, bowoense , is derived from the Tibetan name of the type locality of the new species. We name the new species using the Tibetan name for the type locality in honoring the positive conservation influence of Tibetan culture on the habitats and the inhabiting wildlife at the type locality. We suggest Bowo Mountain Dragon as its English common name, and ñffiżẘ (Pinyin: Bo Wo Long Xi) as its Chinese common name.

Diagnosis. The new species can be diagnosed based on a combination of the following morphological characters: 1) body size small, SVL 45.7–53.7 mm in males, 52.6–54.2mm in females; 2) tail length moderate, TAL 189.1–201.9% SVL in males, 177.3–178.7% in females; 3) HW 65.1–70.5% HL; 4) HLL 72.6–78.6% SVL; 5) nuchal and dorsal crests moderately developed on weak skin folds in males; 6) distinct transverse gular fold present; 7) tympanum concealed; 8) MD 40–44; 9) F4S 15–17; 10) T4S 19–21; 11) PTS 3 or 4; 12) PTY 3–7; 13) PRS 3–6; 14) dorsolateral stripes strongly jagged, Cream Color [Color 12] in males, Sulphur Yellow [Color 80] and faint in females; 15) gular spots present in both sexes, smaller in females, Light Chrome Orange [Color 76] in live males, Dark Spectrum Yellow [Color 78] in live females, absent after preservation; 16) inner lips, oral cavity, and tongue all Light Flesh Color [Color 250]; and 17) distinct vermiculate stripes present on ventral surface of the head.

Comparisons. Previously, the new species had been confused with D. flaviceps , but can be differentiated by having a smaller body size in males (SVL 45.7–53.7mm vs. 68.5–82.1mm), weak skin folds under both dorsal and nuchal crests in males, absence in females (vs. strongly developed and erected in males, presence in females), and by the presence of distinct radial stripes around the eyes (vs. absence), absence of a lateral series of dark, hallow rhomboid patterns between dorsolateral stripes on the dorsum (vs. presence), and by the presence of colorful gular spots in both sexes (vs. absence).

In comparison with species distributed in close proximity and along the same upper Yalong River, D. bowoense sp. nov. differs from D. angustelinea by having a shorter tail (TAL 189.1–201.9% SVL in males, 177.3–178.7% in females vs. 230.2–249.1% in males, 194.30–222.3% in females; Fig. 6 ), better developed nuchal and dorsal crest with weak skin folds in males (vs. feeble crests with no skin folds), and by the presence of distinct dark vermiculate stripes on the ventral surface of the head in both sexes (vs. absence or faint); from D. panchi by having a longer tail (TAL 177.3–178.7% SVL in females vs. 141.8–151.5% in females; Fig. 6 ), and by a distinct gular pattern in females (distinct Dark Spectrum Yellow [Color 78] gular spot vs. mosaic Light Sulphur Yellow [Color 93] patterns); and from D. panlong by having a much smaller body size in males (SVL 45.7–53.7mm vs. 60.2–71.7mm), a concealed tympanum (vs. exposed in most individuals), different coloration of the oral cavity and tongue (Light Flesh Color [Color 250] vs. oral cavity Dark Spectrum Yellow [Color 78], tongue Light Chrome Orange [Color 76]), and by the presence of gular spots in males (vs. absence).

From all three species distributed along the upper Mekong River Valley (i.e. D. drukdaypo , D. iadinum , and D. vela ), D. bowoense sp. nov. can be differentiated from all three species by the presence of a distinct coloration of gular spots (Light Chrome Orange [Color 76] in males, Dark Spectrum Yellow [Color 78] in females vs. absence in D. drukdaypo and D. vela ; Caribbean Blue [Color 168] in males, Medium Greenish Yellow [Color 88] in females of D. iadinum ). Additionally, D. bowoense sp. nov. differs from D. drukdaypo by having distinctively keeled ventral scales of head and body (vs. smooth or feebly keeled) and a much longer tail (TAL 189.1–201.9% SVL in males, 177.3–178.7% in females vs. 153.0–154.4% in males, 132.8–144.0% in females); from D. iadinum by having a distinct body coloration and ornamentation in males (Jet Black [Color 300] to Raw Umber [Color 22] in background, with Cream Color [Color 12], strongly jagged dorsolateral stripes in males vs. Yellowish Spectrum Green [Color 128] to Emerald Green [Color 143] background coloration, with same colored, smooth-edged dorsolateral stripes) and by the absence of multiple regular lateral rows of enlarged scales on the dorsum (vs. presence); and from D. vela by the absence of strongly developed, continuous sail under the vertebral crests in males (vs. feeble).

From all species distributed along the upper Jinsha River ( D. aorun , D. batangense , D. brevicaudum , D. flavilabre , D. qilin , and D. yulongense ), D. bowoense sp. nov. differs by having a distinct coloration of gular spots in both sexes (Light Chrome Orange [Color 76] in males, Dark Spectrum Yellow [Color 78] in females vs. Pale Cyan [Color 157] to Light Caribbean Blue [Color 163] in both sexes of D. aorun ; Pale Cyan [Color 157] in both sexes of D. batangense ; Pale Sulphur Yellow [Color 92] in males only of D. brevicaudum ; Pale Emerald Green [Color 141] to Light Turquoise Green [Color 146] in both sexes of D. flavilabre ; Light Sulphur Yellow [Color 93] in both sexes of D. qilin ; and Chartreuse [Color 89] to Opaline Green [Color 106] in both sexes of D. yulongense ). Additionally, D. bowoense sp. nov. differs from D. aorun by having a shorter tail (189.1–201.9% SVL in males, 177.3–178.7% in females vs. 212.0–221.1% in males, 191.4–207.7% in females); from D. brevicaudum and D. flavilabre by having a longer tail (189.1–201.9% SVL in males, 177.3–178.7% in females vs. 184.1–188.7% in males, 136.8–159.1% in females in D. brevicaudum ; 149.7–179.3% SVL in males, 140.2–152.2% in females in D. flavilabre ) and by distinct inner lip and tongue coloration (Light Flesh Color [Color 250] vs. inner lips Sulphur Yellow [Color 80], tongue Light Orange Yellow [Color 77] in D. brevicaudum ; inner lips Dark Spectrum Yellow [Color 78] in D. flavilabre ); from D. qilin by having a smaller body size in males (SVL 45.7–53.7mm vs. 55.9–66.5mm); and from D. yulongense by having a shorter tail in females (TAL 177.3–178.7% SVL vs. 182.1–211.3%).

Compared to both species distributed along the upper Salween River ( D. laeviventre and D. slowinskii ), D. bowoense sp. nov. differs from D. laeviventre by having fewer MD (MD 40–44 vs. 57–59) and distinctively keeled ventral scales (vs. smooth or feebly keeled); and from D. slowinskii by having concealed tympana (vs. exposed) and a much smaller body size (SVL 45.7–53.7mm in males, 52.6–54.2mm in females vs.> 74mm).

Finally, D. bowoense sp. nov. can be differentiated from D. chapaense , D. fasciatum , D. micangshanense , D. menghaiense , 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 distinct transverse gular fold (vs. absence); from D. dymondi and D. swild by having concealed tympana (vs. exposed); from D. splendidum by having strongly jagged dorsolateral stripes in males (vs. smooth edged) and homogeneous scales on the ventral surface of the head (vs. heterogeneous); from D. hamptoni by having parallel, jagged dorsolateral stripes (vs. diagonally away from midline and smooth); and from D. zhaoermii by having distinct coloration of gular spots in both sexed (Light Chrome Orange [Color 76] in males, Dark Spectrum Yellow [Color 78] in females vs. Chartreuse [Color 89] in males, no gular spots in females) and by having a much smaller body size (SVL 45.7–53.7mm in males, 52.6–54.2mm in females vs. 63.2–81.7mm in males, 61.0– 75.2mm in females).

Description of holotype. Body not dorsally compressed, SVL 53.7mm; tail long, TAL 201.9% SVL; limbs length moderate, FLL 45.2% SVL, HLL 75.6% SVL; head shape moderate, HW 65.1% HL, HD 75.7% HW; SEL 39.5% HL. Rostral rectangular, three times wider than height, two scales away from nasal; nasal oval, somewhat elongated, two scales away from first supralabial; supralabials 10/10, last one longest, weakly keeled; infralabials 10/12, less keeled than supralabials; suborbital scale rows 3/3, each scale bearing distinct, lateral keel; supraciliaries 8/8, overlapping one half of total length with succeeding ones; enlarged, protruding scales between posterior orbit and tympanum, 7/8, each bearing single distinct, lateral keel; tympanum concealed under fine scales; post tympanic conical scales 5/7, moderately developed; enlarged post-rictal scales 3/3, sub-pyramidal in shape. Dorsal head scales heterogeneous in size and shape, all distinctively keeled; single, laterally oriented, Y-shaped ridge present on dorsal snout, starting two scales posterior of rostral to anterior end of orbit; interparietal large, parietal eye distinct; postciliary scale much enlarged, subpyramidal shape; post occipital conical scales moderately differentiated, 3/3.

Dorsal body scales distinctively keeled, heterogeneous in size and shape; axillary scales fine; shoulder fold present, distinct, extending posteriorly over axilla; enlarged scales arranged in two dorsolateral rows on each side of dorsum from neck to pelvis, ones close to dorsal midline, one along mid-dorsolateral stripe; nuchal, dorsal crests well developed on rather weak skin folds, crest scales erected, serrated, differentiated from nearby dorsal sales; middorsal scales 44. Scales of dorsal limbs distinctively keeled, distinctively heterogeneous on hind limbs; Finger IV and Toe IV longest, Finger IV subdigital lamellae 17/16, Toe IV subdigital lamellae 21/21.

Ventral head scales homogeneous in size and shape, distinctively keeled except chin shields; mental pentagonal, completely enclosed by first pair of chin shields; chin shields 7/7, smooth, two scale rows away from infralabials. Gular pouch present, well developed in life, indistinct after preservation; distinct transverse gular fold present, deep. Ventral body and limb scales distinctively keeled, more pronounced than those of ventral head, homogeneous in size and shape, keel of individual scale mostly regularly arranged in lateral rows.

Coloration. The background coloration of the dorsal and lateral surface of the head is Drab Gray [Color 256] to pale Dark Neutral Gray [Color 299]. Four Sepia [Color 286] to Jet Black [Color 300] transverse ornamentation patterns are present on the dorsal surface of head: the first streak is at the posterior edge of the orbit, the second streak is between the anterior edge of the orbit, the third streak is an X-shaped pattern at the midpoint between the eyes and nares, and the last streak is between the nares. Dark Neutral Gray [Color 299] or Jet Black [Color 300] streaks form the radial stripe pattern around the eye, many of which are vague and not clearly defined. A white lip stripe is present in the suborbital region, although the anterior one-third of the stripe is heterogeneous in color and speckled with Medium Neutral Gray [Color 298].

The background coloration of the dorsum is Drab [Color 19] to Dark Neutral Gray [Color 299]. 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. The stripe is strongly jagged, with five rhombus-shaped extensions distributed evenly across the stripe from the shoulder to the pelvis. Dorsolateral stripes are Cream Color [Color 12] with irregular and faded speckles of Olive Horn Color [Color 16]. The nuchal and dorsal crests are Clay Color [Color 18] and distinct from the rest of the dorsal surface coloration. The dorsal surfaces of the limbs and tail are Drag Gray [Color 256], with evenly distributed Dark Neutral Gray [Color 299] transverse bands or streaks running from the proximal to distal ends. The streaks get paler towards the distal ends of the fingers, toes, and the tip of the tail.

The ventral surface of the head is Light Buff [Color 2], with Jet Black [Color 300] short streaks, which are formed by 2–3 rows of black scales. A distinct, Light Chrome Orange [Color 76], triangular-shaped gular spot is present at the center of the gular region. The ventral surfaces of the body, limbs, and tail are uniform Light Buff [Color 2].

Variation. Detailed morphological variation is summarized in Table 4. The paratype male (KIZ 044701) has a shorter tail than the holotype, and the Clay Color [Color 18] of nuchal and dorsal crests are indistinct. Sexual dimorphism is evident despite the small sample size, with males possessing relatively longer tails, relatively shorter trunks, and taller nuchal crests ( Table 4). Furthermore, similar to most Diploderma congeners, D. bowoense sp. nov. are sexually dichromatic, where males have different gular coloration than females. In females, the gular spots are less saturated, and the color is Dark Spectrum Yellow [Color 78]. Between the two females, only one (KIZ 044703) has a similar vertebral stripe as is observed on the male holotype, and the coloration is more saturated (Cinnamon- Rufous [Color 31] to Light Pratt’s Rufous [Color 71]) than the holotype. The other female has six Dark Neutral Gray [Color 299], reverse triangle-shaped blotches scattered evenly along the vertebral line from the neck to the pelvis.

Natural history and conservation. Diploderma bowoense sp. nov. inhabits hot-dry valleys along the middle Yalong River in Muli County, Sichuan Province, China. Currently, the species is known only from the type locality near Bowo Village, but it is likely to be found further northward along the Yalong River Valley. Individuals were found active during the day, basking on rock piles and open ground, but slept on tall bushes at night. Amolops mantzorum , Elaphe carinata , and E. taeniura were found at the type locality of D. bowoense sp. nov.. Although the habitats at the type locality are well protected thanks to the local Tibetan community, the remaining habitat of the species along the main Valley of Yalong River are still threatened by the ongoing development of a nearby hydropower station. We recommend Data Deficient (DD) for the IUCN status of D. bowoense sp. nov. and call for ecological and population studies of the new species in the future.

Figures

FIGURE 3. Comparisons of gular coloration in life between (1) males and (2) females of (A) Diploderma bowoense sp. nov., (B) D. formosgulae sp. nov., (C) D. aorun, (D) D. batangense, (E) D. vela, (F) D. angustelinea, (G) D. qilin, and (H) D. brevicaudum. The smaller inset photos in A1 represent gular coloration of the representation of juvenile male of D. formosgulae sp. nov. Photos by Kai Wang.

FIGURE 6. Boxplot showing the important diagnostic morphometric character, relative tail length, of males and females of the two new species (Diploderma bowoense sp. nov. and D. formosgulae sp. nov.) and their closest congeners. Each box represents the interquartile range (25th to 75th percentile), the thick horizontal bar inside each box indicates the median value, the vertical lines indicate maximum and minimum values. Diploderma panchi and D. panlong are known by a single sex only (female and male, respectively), therefore data of only one sex are available.

FIGURE 7. Dorsolateral view (1) and closeup of the lateral surface of the head (2) of the male holotype KIZ 044700 (A) and female paratype KIZ 044703 (B) of Diploderma bowoense sp. nov. in life. Photos by Kai Wang.

FIGURE 8. Dorsolateral (1) and ventral (2) views of the male holotype KIZ 044700 (A) and female paratype KIZ 044703 (B) of Diploderma bowoense sp. nov. in preservation. Photos by Zhongbing Yu.