Research Article |
Corresponding author: Ying-Yong Wang ( wangyy@mail.sysu.edu.cn ) Academic editor: Angelica Crottini
© 2020 Han Wan, Zhi-Tong Lyu, Shuo Qi, Jian Zhao, Pi-Peng Li, Ying-Yong Wang.
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:
Wan H, Lyu Z-T, Qi S, Zhao J, Li P-P, Wang Y-Y (2020) A new species of the Rana japonica group (Anura, Ranidae, Rana) from China, with a taxonomic proposal for the R. johnsi group. ZooKeys 942: 141-158. https://doi.org/10.3897/zookeys.942.46928
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Rana jiulingensis sp. nov., a new species from Hunan and Jiangxi, southeastern China, is described. The new species is assigned to the R. japonica group. The clade comprising R. jiulingensis sp. nov. and R. dabieshanensis from Anhui is the sister taxon of R. omeimontis from Sichuan. Rana jiulingensis sp. nov. can be distinguished by the significant divergences in the 16S and COI genes, and the combination of following morphological characters: body size medium, SVL 48.3–57.8 mm in adult males and 48.2–57.5 mm in adult females; dorsolateral fold straight; digits without circummarginal grooves; dorsal skin smooth; tibio-tarsal articulation reaching forward beyond the tip of snout; heels overlapping; webbing formula of toes: I 1⅓ – 2 II 1⅓ – 2⅓ III 1½ – 2⅔ IV 3 – 1⅓ V; absence of vocal sacs in males; and presence of creamy white nuptial pad with tiny hoar spines on the finger I and reddish tubercles on loreal and temporal regions in breeding males. Furthermore, based on our results and the previous literature, R. zhengi is synonymized with R. sangzhiensis, and a new species group, the Rana johnsi group, is proposed for the R. johnsi and R. sangzhiensis. Currently, the Rana contains 41 recognized species, and the phylogenetic placements of several species remain unresolved.
morphology, phylogeny, Rana jiulingensis sp. nov., Rana sangzhiensis, Rana zhengi
As the type genus of the family Ranidae Batsch, 1796, the concept of the true-frog genus Rana Linnaeus, 1758 has been discussed for a long time (
Based on morphological comparisons and geographical conditions,
During herpetofaunal surveys in the Luoxiao Range, which is situated between the Jiangxi and Hunan provinces (Fig.
Eight unnamed specimens were collected from multiple localities of Jiangxi and Hunan provinces. All specimens were fixed in 10% buffered formalin, later transferred to 70% ethanol, and deposited in the Museum of Biology, Sun Yat-sen University (
SVL snout–vent length (from tip of snout to posterior margin of vent);
HL head length (from tip of snout to the articulation of the jaw);
HW head width (head width at the commissure of the jaws);
SL snout length (from tip of snout to the anterior corner of the eye);
IN internasal distance (distance between nares);
IO interorbital distance (minimum distance between upper eyelids);
ED eye diameter (from the anterior corner of the eye to posterior corner of the eye);
TD tympanum diameter (horizontal diameter of tympanum);
TED tympanum–eye distance (from anterior edge of tympanum to posterior corner of the eye);
HND hand length (from the proximal border of the outer palmar tubercle to the tip of digit III);
RAD radio-ulna length (from the flexed elbow to the proximal border of the outer palmar tubercle);
FTL foot length (from distal end of shank to the tip of digit IV);
TIB tibial length (from the outer surface of the flexed knee to the heel).
The morphological description follows the consistent definition by
A total of 56 muscular samples of Rana were used, encompassing nine samples of the undescribed specimens, and 47 samples from 12 recognized species. All samples were attained from euthanasia specimens and then preserved in 95% ethanol and stored at –40 °C. Genomic DNA were extracted from muscle tissue samples, using DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. Two mitochondrion genes, namely partial 16S ribosomal RNA gene (16S) and partial cytochrome c oxidase 1 gene (COI), were amplified. Primers used for 16S were L3975 (5'-CGCCTGTTTACCAAAAACAT-3') and H4551 (5'-CCGGTCTGAACTCAGATCACGT-3') following
Localities, voucher information, and GenBank numbers for all samples of the genus Rana used in this study (* = type localities).
ID | Species | Localities | Voucher no. | 16S | COI |
---|---|---|---|---|---|
1 | R. jiulingensis | China: Jiangxi: Mt Guanshan * |
|
MT408985 | MT418647 |
2 | R. jiulingensis | China: Jiangxi: Mt Guanshan * |
|
MT408994 | MT418656 |
3 | R. jiulingensis | China: Jiangxi: Mt Wugong |
|
MT408964 | MT418626 |
4 | R. jiulingensis | China: Jiangxi: Mt Wugong |
|
MT408965 | MT418627 |
5 | R. jiulingensis | China: Hunan: Mt Mufu |
|
MT408984 | MT418646 |
6 | R. jiulingensis | China: Hunan: Mt Dawei |
|
MT408989 | MT418651 |
7 | R. jiulingensis | China: Hunan: Mt Dawei |
|
MT408990 | MT418652 |
8 | R. jiulingensis | China: Hunan: Mt Dawei |
|
MT408991 | MT418653 |
9 | R. jiulingensis | China: Hunan: Mt Dawei |
|
MT408992 | MT418654 |
10 | R. amurensis | China: Heilongjiang: Taiyang Island | SYNU 11100267 | KF020589 | KF020603 |
11 | R. amurensis | China: Liaoning, Zhangwu | SYNU 11100268 | KU343216 | KU343216 |
12 | R. arvalis | Germany: Lower Saxony | No voucher | AY147938 | / |
13 | R. asiatica | China: Xinjiang: 47tuan | KIZ XJ0251 | KX269200 | / |
14 | R. chaochiaoensis | China: Sichuan: Zhaojue * |
|
MT409007 | MT418669 |
15 | R. chaochiaoensis | China: Sichuan: Zhaojue * |
|
MT408957 | MT418619 |
16 | R. chensinensis | China: Shaanxi: Huxian * | KIZ RD05SHX01 | KX269186 | JF939080 |
17 | R. chensinensis | China: Henan: Mt Yawu |
|
MT408962 | MT418624 |
18 | R. chensinensis | China: Henan: Mt Yawu |
|
MT408963 | MT418625 |
19 | R. coreana | South Korea | MMS 223 | KX269202 | MF149928 |
20 | R. coreana | China: Shandong: Mt Kunyu | SYNU 08090641 | MT409004 | MT418666 |
21 | R. culaiensis | China: Shandong: Mt Culai * | KIZ SD080501 | KX269190 | JF939082 |
22 | R. culaiensis | China: Shandong: Mt Culai * | SYNU 08090549 | MT409006 | MT418668 |
23 | R. culaiensis | China: Jiangxi: Mt Wugong |
|
MT408966 | MT418628 |
24 | R. culaiensis | China: Jiangxi: Shanggao |
|
MT408967 | MT418629 |
25 | R. culaiensis | China: Jiangxi: Mt Meiling |
|
MT408971 | MT418633 |
26 | R. dabieshanensis | China: Anhui: Dabie Mountains area * | AHU 2016R001 | MF172963 | / |
27 | R. dybowskii | Russia: Primorye: Khasanskii | MSUZP-IVM-1d | KX269188 | / |
28 | R. dybowskii | China: Jilin: Mt Laoling | SYNU 11070163 | MT409005 | MT418667 |
29 | R. hanluica | China: Hunan: Mt Yangming * |
|
MT408956 | MT418618 |
30 | R. hanluica | China: Hunan: Mt Bamian |
|
MT408969 | MT418631 |
31 | R. hanluica | China: Hunan: Mt Badagong |
|
MT408973 | MT418635 |
32 | R. hanluica | China: Hunan: Mt Yunshan |
|
MT408977 | MT418639 |
33 | R. hanluica | China: Hunan: Mt Xuefeng |
|
MT408999 | MT418661 |
34 | R. hanluica | China: Hunan: Suining |
|
MT409000 | MT418662 |
35 | R. hanluica | China: Hunan: Mt Shunhuang |
|
MT409001 | MT418663 |
36 | R. hanluica | China: Guizhou: Mt Leigong |
|
MT408959 | MT418621 |
37 | R. hanluica | China: Guizhou: Mt Fanjing |
|
MT408976 | MT418638 |
38 | R. hanluica | China: Jiangxi: Mt Jinggang |
|
MT408968 | MT418630 |
39 | R. hanluica | China: Jiangxi: Mt Qiyun |
|
MT408970 | MT418632 |
40 | R. hanluica | China: Guangxi: Longsheng |
|
MT408960 | MT418622 |
41 | R. hanluica | China: Guangxi: Mt Dupangling |
|
MT408980 | MT418642 |
42 | R. hanluica | China: Guangdong: Renhua |
|
MT408998 | MT418660 |
43 | R. huanrenensis | China: Liaoning: Huanren * | SYNU 07040035 | KF204642 | KX139725 |
44 | R. huanrenensis | China: Liaoning: Huanren * | y-d20130058 | KT588071 | KT588071 |
45 | R. japonica | Japan: Isumi-shi: Chiba Prefecture | KIZ YPX11775 | KX269220 | JF939101 |
46 | R. japonica | Japan: Isumi-shi: Chiba Prefecture | NNRj | AB728192 | / |
47 | R. jiemuxiensis | China: Hunan: Jiemuxi * |
|
MT408975 | MT418637 |
48 | R. jiemuxiensis | China: Hunan: Jiemuxi * |
|
MT409008 | MT418670 |
49 | R. johnsi | Vietnam: Lam Dong: Loc Bao | ABV 00203 | KX269182 | / |
50 | R. kukunoris | China: Qinghai: Qinghai Lake * | KIZ CJ06102001 | KX269185 | JF939073 |
51 | R. kukunoris | China: Sichuan: Hongyuan |
|
MT409009 | MT418671 |
52 | R. kukunoris | China: Sichuan: Hongyuan |
|
MT408993 | MT418655 |
53 | R. longicrus | China: Taiwan:Taipei * | Not given | AB058881 | / |
54 | R. longicrus | China: Taiwan: Miaoli: Xiangtianhu | NMNS 15022 | KX269189 | / |
55 | R. longicrus | China: Fujian: Mt Yashu |
|
MT408987 | MT418649 |
56 | R. longicrus | China: Jiangxi: Mt Magu |
|
MT408996 | MT418658 |
57 | R. longicrus | China: Jiangxi: Mt Qiyun |
|
MT408961 | MT418623 |
58 | R. longicrus | China: Jiangxi: Mt Jiulian |
|
MT408978 | MT418640 |
59 | R. longicrus | China: Jiangxi: Mt Sanbai |
|
MT408986 | MT418648 |
60 | R. longicrus | China: Jiangxi: Suichuan |
|
MT408997 | MT418659 |
61 | R. longicrus | China: Guangdong: Renhua |
|
MT408954 | MT418616 |
62 | R. longicrus | China: Guangdong: Mt Nankun |
|
MT408955 | MT418617 |
63 | R. longicrus | China: Guangdong: Pu’ning |
|
MT408979 | MT418641 |
64 | R. longicrus | China: Guangdong: Mt Tonggu |
|
MT408981 | MT418643 |
65 | R. longicrus | China: Guangdong: Yingde |
|
MT409003 | MT418665 |
66 | R. maoershanensis | China: Guangxi: Mt Maoershan * | SYNU 08030061 | HQ228162 | / |
67 | R. maoershanensis | China: Guangxi: Mt Maoershan * | SYNU 08030062 | HQ228163 | / |
68 | R. luanchuanensis | China: Henan: Luanchuan * | KIZ 047452 | / | MF149923 |
69 | R. luanchuanensis | China: Henan: Luanchuan * | KIZ 047393 | / | MF149924 |
70 | R. omeimontis | China: Sichuan: Mt Emei * |
|
MT408982 | MT418644 |
71 | R. omeimontis | China: Sichuan: Mt Emei * |
|
MT408983 | MT418645 |
72 | R. sangzhiensis | China: Hunan: Mt Tianping * |
|
MT408972 | MT418634 |
73 | R. sangzhiensis | China: Hunan: Mt Tianping * |
|
MT408974 | MT418636 |
74 | R. zhengi | China: Sichuan: Hongya: Zhangcun * | SCUM 0405190CJ | KX269206 | MF149929 |
75 | R. zhengi | China: Sichuan: Hongya: Zhangcun * | KIZ YP06057 | DQ289104 | / |
76 | R. sauteri | China: Taiwan: Kaohsiung * | SCUM 0405175CJ | KX269204 | / |
77 | R. shuchinae | China: Sichuan: Zhaojue |
|
KX269210 | / |
78 | R. zhenhaiensis | China: Zhejiang: Hangzhou | SYNU 08040100 | KF020599 | KF020613 |
79 | R. zhenhaiensis | China: Zhejiang: Zhenhai * | KIZ 0803271 | KX269218 | JF939065 |
80 | R. zhenhaiensis | China: Zhejiang: Fenghua |
|
MT408988 | MT418650 |
81 | R. zhenhaiensis | China: Jiangxi: Mt Tongbo |
|
MT408958 | MT418620 |
82 | R. zhenhaiensis | China: Jiangxi: Mt Guanshan |
|
MT408995 | MT418657 |
83 | R. zhenhaiensis | China: Jiangxi: Mt Yangjifeng |
|
MT409002 | MT418664 |
For phylogenetic analyses, 26 additional sequences from all known Chinese congeners of the subgenusRana (except R. (R.) chevronta) and an out-group sequence of R. (Liuhurana) shuchinae Liu, 1950 were obtained from GenBank and incorporated into our dataset. Detailed information of these materials is shown in Table
The unnamed specimens from Jiangxi and Hunan are assigned to the Rana japonica group based on the following combined characteristics: digits without circummarginal grooves, and dorsolateral fold distinct, extending straight from the posterior margin of the upper eyelid to above the groin. Therefore, we compare the new species with the species of the R. japonica group.
The new species differs from Rana dabieshanensis in the following characters: head length significantly larger than head width, HW/HL 0.82 in males and 0.85 in females (vs almost equal); supratympanic fold absent (vs distinct); tympanum diameter significantly smaller than eye diameter with TD/ED = 0.63–0.87 (vs equal); relative toe lengths I < II < III < V < IV (vs I < II < V < III < IV); toe webbing formula I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V (vs I 2 – 1 II 2+ – 1+ III 3 – 2 IV 2 – 2+ V); and nuptial pad creamy white in breeding males (vs gray-blackish).
The new species differs from R. omeimontis as follows: body size smaller, SVL = 48.2–57.5 mm in adult females (vs 61.7–70.3 mm in females); head length significantly larger than head width, HW/HL = 0.82 in males and 0.85 in females (vs head length slightly larger than head width, HW/HL = 0.94 in males and 0.92 in females); and supernumerary tubercles present below the bases of each finger (vs absent).
The new species further differs from R. hanluica as follows: supratympanic fold absent (vs present); toe webbing formula I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V (vs I 1⅓ – 1⅔ II 1 – 2 III 1⅓ – 2½ IV 2⅓ – 1 V); reddish tubercles present on loreal and temporal regions in breeding males (vs absent, but white horny spines present around loreal and temporal regions, upper eyelids, and snout in breeding males). The new species differs from R. longicrus in having: internarial distances larger than interorbital distances (vs smaller) and toe webbing formula I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V (vs I 1⅔ – 2⅓ II 1½ – 2⅔ III 1⅔ – 3½ IV 3⅓ – 1½ V); from R. zhenhaiensis: supratympanic fold absent (vs present), dorsolateral fold extending straight from posterior margin of upper eyelid to above groin (vs dorsolateral fold slightly curved above tympanum), two outer metacarpal tubercles distinctly separated (vs merged at base), tibio-tarsal articulation reaching forward beyond tip of snout (vs around nostril), and nuptial pad creamy white in breeding males (vs gray or gray-brownish); from R. culaiensis: dorsolateral fold extending straight from posterior margin of upper eyelid to above groin (vs dorsolateral fold slightly curved above tympanum), and tibio-tarsal articulation reaching forward beyond tip of snout (vs at nostril); from R. jiemuxiensis: dorsolateral fold extending straight from posterior margin of upper eyelid to above groin (vs dorsolateral fold slightly curved above tympanum), head length significantly larger than head width (vs slightly larger), internarial distances larger than interorbital distances (vs smaller), and two outer metacarpal tubercles distinctly separated (vs merged at base); from R. chaochiaoensis: supratympanic fold absent (vs present), internarial distances larger than interorbital distances (vs smaller), and toe webbing formula I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V (vs I 1 – 1⅔ II 1⅓ – 2 III 1½ – 2½ IV 2⅔ – 1 V); from R. japonica: outer metacarpal tubercles present (vs absent), tibio-tarsal articulation reaching forward beyond tip of snout (vs reaching or beyond tip of snout in males, reaching at center of eye or beyond nostril in females), nuptial pad creamy white and divided into three parts (vs nuptial pads grayish brown or yellowish brown and divided into two parts).
From Rana chevronta, which lacks molecular data, the new species can be distinguished by its larger body size, SVL = 48.3–57.8 mm in adult males (vs 39.7–44.0 mm), head length significantly larger than head width (vs almost equal), relative finger lengths I < II < IV < III (vs II < IV < I < III), and nuptial pad creamy white and divided into three parts in breeding males (vs purplish gray and undivided).
The ML and BI analyses resulted in essentially identical topologies and are integrated in Figure
The Rana samples representing the new species are grouped in a distinct and robust monophyletic lineage with high support (BPP = 1.00 and BS = 100) and low divergence (mean 0.3%, ranging 0.0–0.6% in COI, and mean 0.1%, ranging 0.0–0.5% in 16S); they form a separate evolutionary lineage within the R. japonica group. This lineage from Jiangxi and Hunan is close to R. dabieshanensis from Anhui and R. omeimontis from Sichuan. The smallest genetic distance between this lineage and a previously recognized species is 3.4–4.0% in COI (with R. omeimontis) and 1.6–2.0% in 16S (with R. dabieshanensis), which are significant when compared to all other recognized species (e.g. 2.8–3.6% in COI between R. longicrus and R. culaiensis; 1.2–1.3% in 16S between R. dabieshanensis and R. omeimontis).
Therefore, based on the significant morphological differences and phylogenetic divergence, these specimens from Jiangxi and Hunan represent a distinct evolutionary lineage and are described as a new species, Rana jiulingensis sp. nov.
Seven adult specimens. Females
The specific name jiulingensis is in reference to the type locality, Guanshan Nature Reserve in Jiuling Mountains.
Jiuling Mountains Brown Frog (in English), Jiu Ling Shan Lin Wa (九岭山林蛙 in Chinese)
Rana jiulingensis sp. nov. is distinguished by the following morphological characteristics: (1) body medium-sized, SVL = 48.3–57.8 (51.7 ± 4.3, n = 4) mm in adult males, 48.2–57.5 (50.8 ± 4.4, n = 4) mm in adult females; (2) head length significantly larger than head width; (3) supratympanic fold absent; (4) dorsolateral fold distinct and thin, extending straight from posterior margin of upper eyelid to above groin; (5) internarial distances larger than interorbital distances; (6) tympanum diameter significantly smaller than eye diameter, TD/ED = 0.63–0.87; (7) fingers without circummarginal grooves, unwebbed, relative finger lengths I < II < IV < III; (8) presence of supernumerary tubercles below the bases of each finger, presence of three separated metacarpal tubercles; (9) toes without circummarginal grooves, toe webbing formula: I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V, relative toe lengths I < II < III < V < IV; (10) tibio-tarsal articulation reaching forward beyond tip of snout; (11) heels overlapping; (12) dorsal skin smooth, flanks smooth with few granules; (13) absence of vocal sacs in males; (14) breeding males possess creamy white nuptial pad with tiny hoar spines on the finger I, divided into three parts; (15) presence of reddish tubercles on loreal and temporal regions in breeding males.
Forearms 0.19 of SVL and hand 0.26 of SVL; fingers slender, without web but with narrow fringe; tip of fingers rounded, not expanded, without circummarginal grooves; relative finger lengths I < II < IV < III; subarticular tubercles significantly prominent, rounded; distinct, small, rounded supernumerary tubercles below the bases of each finger; inner metacarpal tubercle indistinct, ovoid, partly covered by nuptial pad; two outer metacarpal tubercles distinctly separated, slightly larger, long elliptic; nuptial pad with tiny spines on the finger I, divided into three parts, the basal one around the inner metacarpal tubercle and partly covering it, the largest one from the edge of the basal one to the subarticular tubercle of finger I, the smallest one extending from the edge of the biggest one to the tip of finger I.
Tibia 0.63 of SVL and foot 0.88 of SVL; heels overlapping when hindlimbs flexed at right angles to axis of body; tibio-tarsal articulation reaching forward beyond the tip of snout when hindlimb stretched along the side of the body; relative toe lengths I < II < III < V < IV; toes webbing formula: I 1⅓ – 2 II 1⅓ – 2⅓ III 1 ½ – 2⅔ IV 3 – 1⅓ V; absence of lateral fringes on the lateral edges of toes I and V; subarticular tubercles oval and distinct; inner metatarsal tubercle large, ovoid, outer metatarsal tubercle small.
Dorsal skin smooth with sparse tiny granules; several small tubercles on flank; supratympanic fold absent; dorsolateral fold distinct and thin, extending straight from posterior margin of upper eyelid to above groin; several tiny granules on the skin of loreal and temporal regions; ventral surface smooth, large flattened tubercles densely arranged on the rear of thigh and around vent.
In life, dorsal surface yellowish brown with few black spots; black speckles forming a linear stripe between eyelids; dorsolateral fold intermittently edged with black on two sides; loreal region yellowish; temporal region yellowish, slightly tinged with grey; tiny granules on loreal and temporal regions reddish; dorsal forelimbs and hindlimbs reddish with indistinct greenish grey transverse bars. Throat yellowish; chest and belly creamy white; ventral surface of forelimbs and hindlimbs flesh color; nuptial pad creamy white; tubercles around vent yellowish.
In preservative, dorsal surface turns grey with black spots and light grey patches; limbs taupe with brown transverse bars. Ventral surface white, with greyish mottling on throat and belly; ventral surface of limbs beige; hands and toe webs dark grey.
Measurements of type series specimens are given in Table
Measurements (in mm) of the type series of Rana jiulingensis sp. nov. (* = holotype).
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|
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Sex | Male | Male | Male | Male | Female | Female | Female | Female |
SVL | 57.8 | 51.6 | 48.3 | 49.1 | 57.5 | 48.4 | 49.4 | 48.2 |
HL | 21.6 | 19.3 | 18.4 | 17.7 | 22.3 | 18.9 | 18.2 | 19.4 |
HW | 18.4 | 17.0 | 15.7 | 12.6 | 19.3 | 16.1 | 15.5 | 15.8 |
SL | 7.8 | 7.5 | 7.3 | 7.5 | 8.1 | 7.1 | 7.2 | 7.4 |
IN | 4.1 | 3.8 | 3.5 | 3.2 | 4.2 | 3.6 | 4.1 | 4.1 |
IO | 3.4 | 2.7 | 3.0 | 3.1 | 3.6 | 3.4 | 3.3 | 3.3 |
ED | 6.3 | 5.5 | 5.2 | 5.0 | 5.2 | 4.4 | 4.7 | 4.6 |
TD | 4.6 | 3.5 | 3.3 | 3.9 | 4.5 | 3.7 | 3.2 | 3.4 |
TED | 1.9 | 2.0 | 1.7 | 2.0 | 1.7 | 1.7 | 1.7 | 1.6 |
HND | 15.0 | 14.0 | 14.0 | 12.5 | 15.3 | 13.9 | 13.2 | 12.5 |
RAD | 11.2 | 11.2 | 9.0 | 9.3 | 10.5 | 10.4 | 10.9 | 10.1 |
FTL | 50.7 | 44.2 | 41.7 | 42.8 | 47.6 | 43.8 | 44.3 | 41.1 |
TIB | 36.6 | 30.4 | 29.5 | 30.7 | 36.1 | 31.8 | 31.8 | 29.0 |
Currently, Rana jiulingensis sp. nov. is known from Guanshan Nature Reserve in the Jiuling Mountains and Mount Wugong in the Wugong Mountains of northwestern Jiangxi, and Mount Mufu and Mount Dawei in the Mufu Mountains of northeastern Hunan. This suggests that its geographic distribution is the central and northern parts of the Luoxiao Range (Fig.
All recognized species of the subgenusRana from China (except for R. chevronta) are included in our work for morphological and molecular analyses. Four monophyletic clades are supported by high values (BPP = 1.00 and BS > 85, respectively) in the phylogenetic tree. Three of them correspond to the morphologically recognized R. japonica group, R. chensinensis group, and R. amurensis group. The fourth, unnamed monophyletic clade includes R. johnsi, R. sangzhiensis, and R. zhengi. Within this unnamed clade, R. sangzhiensis and R. zhengi cluster together with significant support (BPP = 1.00 and BS = 100) and little divergence (0.0–0.4% in COI and 0.0–0.4% in 16S), which is consistent with the original morphological identification by
Within the Rana japonica group, the genetic divergences among three species, R. longicrus, R. zhenhaiensis, and R. culaiensis, are relatively closer than other species. Additionally, the validations of these species have been supported by the morphological examinations (
The discovery of Rana jiulingensis sp. nov. increases the diversity of the genus Rana in the Luoxiao Range to five species (Fig.
We thank the Hunan Badagongshan National Nature Reserve, Fujian Yashushan Nature Reserve, Jian Wang, Chun-Lin Liao, Hai-Long He, Zhao-Chi Zeng, Yu-Long Li, and Zheng-Yan Zhou for their help with the fieldwork. We thank Zhen-Hua Liu, Zi-Chen Zhou, Yang Chen, and Robert Forsyth for their help in polishing the manuscript. We thank Angelica Crottini and two anonymous reviewers for their helpful suggestions on our work. This work was supported by the Project of Comprehensive Scientific Survey of Luoxiao Mountains Region of Ministry of Science and Technology, China (no. 2013FY111500), the Project of Scientific Investigation on the Amphibian, Reptilian and Avian Animals in Jiangxi Jiulianshan National Nature Reserve, the Project of Animal Diversity Survey and Monitoring System Construction of Guangdong Shimentai National Nature Reserve, and the Project of Survey of Terrestrial Vertebrate Diversity in Guangdong Danxiashan National Nature Reserve.
Specimens examined
Rana chaochiaoensis (3): China: Sichuan: Zhaojue County (type locality):
Rana chensinensis (2): China: Henan: Mt Yawu:
Rana culaiensis (4): China: Jiangxi: Mt wugong:
Rana hanluica (35): China: Hunan: Mt Yangming (type locality):
Rana jiemuxiensis (2): China: Hunan: Jiemuxi Nature Reserve (tpye locality):
Rana kukunoris (7): China: Sichuan: Hongyuan County:
Rana longicrus (18): China: Fujian: Mt Yashu:
Rana omeimontis (5): China: Sichuan: Mt Emei (type locality):
Rana zhenhaiensis (7): China: Zhejiang: Fenghua District:
Tables S1, S2
Data type: Tables for genetic distances
Explanation note: Table S1. Pairwise distances based on COI gene among all sample used in this study. Table S2. Pairwise distances based on 16S gene among all sample used in this study.