Odorrana damingshanensis, Chen & Mo & Lin & Qin, 2024

Chen, Wei-Cai, Mo, Yun-Ming, Lin, Li & Qin, Kun, 2024, A new species of Odorrana Fei, Ye & Huang, 1990 (Amphibia, Anura, Ranidae) from central Guangxi, China with a discussion of the taxonomy of Odorrana (Bamburana), ZooKeys 1190, pp. 131-152 : 131

Odorrana damingshanensis sp. nov.

Figs 3 , 4

Type material examined.

Holotype. NNU 00690, adult male, from the Damingshan National Nature Reserve, Wuming District, Nanning City, Guangxi, China (23.4637°N, 108.4869°E; elevation 1159 m), collected by Weicai Chen on 25 April 2022. Paratypes. NNU 00689 and NNU 00691, adult males; NNU 00692 and NNU 00693, adult females, collected at the same site and time as the holotype.

Diagnosis.

Based on both molecular analyses and specific morphological traits, these specimens were assigned to the genus Odorrana . The distinguishing morphological characteristics of these species include dilated and tapering tips of the digits, disks with circummarginal grooves and a longer vertical diameter than horizontal diameter, fully webbed toes, the absence of a tarsal fold, a thick first finger with a distinct nuptial pad, sawtooth spinules on the upper lip, and well-defined dorsolateral folds ( Fei et al. 2001, 2005).

Odorrana damingshanensis sp. nov. can be distinguished from other species in its genus by the following combination of characters: (1) medium body size (SVL 52.3-54.8 mm in males and 74.8-81.2 mm in females); (2) SVL of female/SVL of male = 1.46; (3) sawtooth spinules on the upper lip; (4) snout obtusely rounded without significantly extending beyond the lower jaw; (5) well-defined dorsolateral folds; (6) horny tubercles on the rear of the back; (7) presence of outer metatarsal tubercles; (8) absence of a supratympanic fold; (9) highly dilated nuptial pad with velvety spinules on the dorsolateral surface of Finger I; (10) distinct expansion of the tips of the second, third, and fourth fingers, with the first slightly enlarged; expanded tips of the toes with distinct circummarginal grooves; (11) distinct maxillary gland with tiny spines; and (12) external lateral vocal sac (Fig. 3A-I ).

Description of holotype.

Head longer than wide (HDL/HDW = 1.23); snout obtusely rounded in dorsal view, but not strongly projecting beyond margin of lower jaw (Fig. 3G ); canthus rostral distinct, loreal region concave; nostrils oval, oblique, and closer to tip of snout than eye; upper lip with sawtooth spinules (Fig. 3G ); eye diameter less than snout length (EYE/SNT = 0.88); interorbital region flat with a pineal; interorbital distance less than eye diameter (IOD/EYE = 0.75); internostril distance less than eye diameter (IN/EYE = 0.88); tympanum distinct, rounded, 57% eye diameter, slightly concave relative to skin of temporal region; supratympanic fold absent; vomerine teeth on two oblique ridges, closed each other than choana; tongue elongated, deeply notched posteriorly; pupil horizontally oval; and external lateral vocal sac (Fig. 3I ).

Forelimbs stout, relative length of fingers II <IV <I <III; tips of the second, third and fourth distinctly expanded, but the first slightly enlarged; tips of all fingers with circummarginal grooves; Finger III disk width less than tympanum diameter (FD3/TMP = 0.62); finger webbing absent; subarticular tubercles prominent, rounded, formula 1, 1, 2, 2; inner and outer palmar tubercles distinct; nuptial pad on lateral surface of Finger I strongly dilated with velvety spinules, extending from hand base to level of subarticular tubercle (Fig. 3E, F ). Tips of toes expanded, with distinct circummarginal grooves; relative length of toes I <II <III ≈ V <IV; toes entirely webbed; subarticular tubercles distinct, rounded, formula 1, 1, 2, 3, 2; inner metatarsal tubercle elongated; outer metatarsal tubercle conical (Fig. 3H ).

Body surface shagreened; rear of the back with horny tubercles; ventral surface of venter, forelimbs, and thighs smooth; flanks shagreened; dorsal of forelimbs and hindlimbs shagreened, and hindlimbs with sparse tubercles; two distinct maxillary glands with tiny spinules (Fig. 3I ).

Coloration in life.

Dorsum grey-beige, with irregular grass-green blotches; a discrete darker brown stripe from tip of snout, across canthus rostral, along the inferior dorsolateral fold, finally ending at the anterior of groin; pineal gland grass-green; cream white stripe from anterior of upper lip to maxillary glands; tympanic region brown, some creamy white tubercles around the tympanum; the upper part of belly, chest, and throat with irregular grey cloud, but the lower part of belly creamy white without spots; ventral surface of thighs and forelimbs incarnadine without spots; forelimbs and hindlimbs with pale brown crossbars, three on lower arm, four on thigh and four on tibia; pupil black with orange border; iris creamy yellow, but the posterior iris pale jacinth; and velvety nuptial pad creamy white (Fig. 3A-I ).

Coloration in preservative.

Dorsum brown; bars on forelimbs and hindlimbs darker brown; horny tubercles on the rear of the back turned into creamy white; creamy white nuptial pad turned into grey; the upper part of belly, throat, and chest with brown cloud; the lower part of belly immaculate creamy white; the ventral surface of the hindlimbs creamy yellow; external lateral vocal sac pale green and projecting distinctly (Fig. 3I ).

Etymology.

The specific name of this species, Odorrana damingshanensis , is derived from its discovery locality, Damingshan National Nature Reserve. In English, it is suggested to be called the Damingshan Bamboo-leaf Frog. In Chinese, it is known as 大明山竹叶蛙(Dà Míng Shān Zhú Yè Wā).

Distribution and ecology.

Odorrana damingshanensis sp. nov. is a newly discovered species found in the Damingshan National Nature Reserve, located in Wuming District, Nanning City, Guangxi, China. This species was observed near slow-flowing rocky streams, which were ~ 2-3 m wide and 10-30 cm deep (Fig. 1B ). The frogs were often seen sitting on rocks, and interestingly, one specimen (NNU 00691) was found on a leaf near a stream, while another specimen (NNU 00693) was perched on a dry branch above a stream. The surrounding vegetation in the area consists of evergreen forest, and the elevation is ~ 1200 m with an ambient temperature of 20 °C in April. During the survey, no advertisement calls were heard; however, both female specimens were gravid, carrying creamy yellow eggs without black poles (Fig. 4A ). Additionally, amplexus behavior was observed when males encountered females indoors (Fig. 4B ). The breeding season of O. damingshanensis sp. nov. is speculated to occur between April and May. Other sympatric species in the area include Gracixalus jinxiuensis and Quasipaa shini .

Sexual dimorphism and variation.

The measurements of O. damingshanensis sp. nov. are provided in Table 3 . Females of this species were observed to be significantly larger than males in terms of SVL, with a ratio of 1.46. The specimens show variation in dorsal colors: NNU 00691 displays a grass green coloration with brown blotches (Fig. 4C ), while NNU 00689 exhibits a pale beige coloration with grass green blotches (Fig. 4D ). Males have a higher density of horny tubercles on the rear of their backs compared to females (Fig. 4C-F ).

Comparisons.

Odorrana damingshanensis sp. nov. shares morphological similarities with closely phylogenetically related species, including O. exiliversabilis , O. nasica , O. nasuta , O. tormota , O. trankieni , O. versabilis , and O. yentuensis (Fig. 4G-J , Table 2 ). These similarities can be observed in its protruding snout, slender limbs, sawtooth spinules on the upper lip, and distinct dorsolateral folds. However, O. damingshanensis sp. nov. can be distinguished from O. exiliversabilis by the noticeably larger body size in females (SVL 74.8-81.2 mm vs SVL 51.8-61.8 mm); presence of horny tubercles on the rear of the back (vs absence); and presence of an external lateral vocal sac (vs an internal subgular vocal sac) ( Fei et al. 2001, 2012; Li et al. 2001). Odorrana damingshanensis sp. nov. differs from O. nasica in having a relatively larger body size in males (SVL 52.3-54.8 mm vs SVL 41.0-46.0 mm); absence of strongly protruding snout (vs presence); a distinct pineal body (vs invisible); an elongated inner metatarsal tubercle (vs an oval inner metatarsal tubercle); absence of whitish spinules scattered ventrally near the groin (vs presence of such spinules) ( Yang 1991; Zhang and Wen 2000; Stuart and Chan-ard 2005; Tran et al. 2008). Odorrana damingshanensis sp. nov. differs from O. nasuta in having a relatively smaller body size in males (SVL 52.3-54.8 mm vs SVL 57.1-63.2 mm); SVL of female/SVL of male = 1.46 (vs the ratio of 1.24), distinct maxillary gland with tiny spines (vs absent tiny spines on maxillary gland); presence of horny tubercles on the rear of the back (vs smooth); absence of strongly protruding snout (vs presence) ( Fei et al. 2001, 2012; Li et al. 2001). Odorrana damingshanensis sp. nov. differs from O. tormota by the absence of a deeply sunk tympanum forming an external auditory canal (vs presence of a deeply sunk tympanum); conspicuously larger body size (SVL 52.3-54.8 mm in males and 74.8-81.2 mm in females vs SVL 32.0-36.3 mm in males and 59.30-60.0 mm in females); presence of sawtooth spinules on upper lip (vs absence) ( Wu 1977; Fei et al. 2012; AmphibiaChina 2023). Odorrana damingshanensis sp. nov. differs from O. trankieni in having a conspicuously smaller body size (SVL 52.3-54.8 mm in males and 74.8-81.2 mm in females vs SVL 75.2-84.1 mm in males and 86.8-95.9 mm in females); SVL of female/SVL of male = 1.46 (vs similar body sizes for males and females); absence of strongly protruding snout (vs presence); presence of horny tubercles on the rear of the back (vs smooth); relative lengths of fingers II <IV <I <III (vs IV <II <I <III); distinct maxillary gland with tiny spines (vs absent tiny spines)( Orlov et al. 2003; Pham et al. 2020). Odorrana damingshanensis sp. nov. differs from O. versabilis in having a conspicuously smaller body size in males (SVL 52.3-54.8 mm vs SVL 70.4-77.2 mm); presence of horny tubercles on the rear of the back (vs smooth); distinct maxillary gland with tiny spines (vs absent tiny spines); external lateral vocal sac (vs internal subgular vocal sac) ( Liu and Hu 1962; Fei et al. 2001, 2012; Li et al. 2001). Odorrana damingshanensis sp. nov. differs from O. yentuensis in having a notably larger body size (SVL 52.3-54.8 mm in males and 74.8-81.2 mm in females vs SVL 41.7-46.2 mm in males and 59.3-65.7 mm in females); shagreened body surface (vs smooth); distinct maxillary gland with tiny spines (vs absent tiny spines on maxillary gland); irregular grey cloud on the upper part of belly, chest, and throat, but creamy white the lower part of belly without spots (vs yellowish white ventral side of body without spots)( Tran et al. 2008; Lu et al. 2016).

Finally, O. damingshanensis sp. nov. can be distinguished from other Odorrana species by the presence of sawtooth spinules on the upper lip (vs absent sawtooth spinules on the upper lip, O. absita , O. amamiensis , O. andersonii , O. anlungensis , O. arunachalensis , O. aureola , O. bacboensis , O. banaorum , O. bolavensis , O. cangyuanensis , O. chapaensis , O. chloronota , O. concelata , O. dulongensis , O. fengkaiensis , O. geminata , O. gigatympana , O. grahami , O. graminea , O. hainanensis , O. heatwolei , O. hejiangensis , O. hosii , O. huanggangensis , O. ichangensis , O. indeprensa , O. ishikawae , O. jingdongensis , O. junlianensis , O. khalam , O. kuangwuensis , O. kweichowensis , O. leporipes , O. liboensis , O. lipuensis , O. livida , O. lungshengensis , O. macrotympana , O. margaretae , O. mawphlangensis , O. monjerai , O. morafkai , O. mutschmanni , O. nanjiangensis , O. narina , O. orba , O. sangzhiensis , O. schmackeri , O. sinica , O. splendida , O. supranarina , O. swinhoana , O. tianmuii , O. tiannanensis , O. tormota , O. utsunomiyaorum , O. wuchuanensis , and O. yizhangensis ); the presence of external lateral vocal sac (vs absent vocal sac, O. arunachalensis , O. concelata , O. heatwolei , O. hosii , O. ichangensis , O. kuangwuensis , O. leporipes , O. liboensis , O. lipuensis , O. livida , O. margaretae , O. mawphlangensis , O. monjerai , O. mutschmanni , O. narina , O. sangzhiensis , O. schmackeri , O. splendida , O. supranarina , O. wuchuanensis ); well-defined dorsolateral folds (vs absent dorsolateral folds, O. andersonii , O. anlungensis , O. arunachalensis , O. aureola , O. bacboensis , O. cangyuanensis , O. chapaensis , O. chloronota , O. concelata , O. dulongensis , O. fengkaiensis , O. geminata , O. grahami , O. hainanensis , O. heatwolei , O. hejiangensis , O. huanggangensis , O. ichangensis , O. ishikawae , O. jingdongensis , O. junlianensis , O. kuangwuensis , O. kweichowensis , O. liboensis , O. lipuensis , O. lungshengensis , O. macrotympana , O. margaretae , O. mawphlangensis , O. morafkai , O. mutschmanni , O. nanjiangensis , O. sangzhiensis , O. schmackeri , O. sinica , O. splendida , O. swinhoana , O. tianmuii , O. tiannanensis , O. wuchuanensis , and O. yizhangensis ); horny tubercles on the rear of the back (vs absent horny tubercles on the rear of the back, O. absita , O. amamiensis , O. andersonii , O. anlungensis , O. arunachalensis , O. aureola , O. bacboensis , O. banaorum , O. bolavensis , O. cangyuanensis , O. chapaensis , O. chloronota , O. concelata , O. dulongensis , O. fengkaiensis , O. geminata , O. gigatympana , O. grahami , O. graminea , O. hainanensis , O. heatwolei , O. hejiangensis , O. hosii , O. huanggangensis , O. ichangensis , O. indeprensa , O. ishikawae , O. jingdongensis , O. junlianensis , O. khalam , O. kuangwuensis , O. kweichowensis , O. leporipes , O. liboensis , O. lipuensis , O. livida , O. lungshengensis , O. macrotympana , O. margaretae , O. mawphlangensis , O. monjerai , O. morafkai , O. mutschmanni , O. nanjiangensis , O. narina , O. orba , O. sangzhiensis , O. schmackeri , O. sinica , O. splendida , O. supranarina , O. swinhoana , O. tianmuii , O. tiannanensis , O. utsunomiyaorum , O. wuchuanensis , and O. yizhangensis ).

Figures

Figure 1. A distributions of the subgenus Odorrana (Bamburana), and B habitat of O. (B.) damingshanensis sp. nov. Abbreviations: DMS = Damingshan, DYS = Dayaoshan, SWDS = Shiwandashan, HP = Huaping, DD = Diding.

Figure 3. The holotype of O. (B.) damingshanensis sp. nov. (NNU 00690) A dorsal view B ventral view C dorsolateral view D rear of the back with horny tubercles and dorsal view of thighs E ventral view of hand F nuptial pad with velvety spinules G ventral view of snout H ventral view of foot, and I external lateral vocal sac and tiny spines on maxillary glands.

Figure 4. A female with creamy yellow eggs without pigmented poles B amplexus C dorsal view of NNU 00691 D dorsal view of NNU 00689 E dorsal view of NNU 00692 F dorsal view of NNU 00693 G dorsolateral view of O. (B.) yentuensis (NHMG 1401036, adult male) H dorsolateral view of O. (B.) versabilis (NNU 00638, adult male) I dorsal view of O. (B.) nasica (NNU 00663, adult female) J dorsolateral view of O. (B.) trankieni (NHMG 141107, adult male).