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Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China

Yao Li Dan-Dan Zhang Zhi-Tong Lyu Jian Wang Yu-Long Li Zu-Yao Liu Hong-Hui Chen Ding-Qi Rao Zhi-Fang Jin Chang-You Zhang Ying-Yong Wang

Yao Li, Dan-Dan Zhang, Zhi-Tong Lyu, Jian Wang, Yu-Long Li, Zu-Yao Liu, Hong-Hui Chen, Ding-Qi Rao, Zhi-Fang Jin, Chang-You Zhang, Ying-Yong Wang. Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China. Zoological Research, 2020, 41(2): 105-122. doi: 10.24272/j.issn.2095-8137.2020.033
Citation: Yao Li, Dan-Dan Zhang, Zhi-Tong Lyu, Jian Wang, Yu-Long Li, Zu-Yao Liu, Hong-Hui Chen, Ding-Qi Rao, Zhi-Fang Jin, Chang-You Zhang, Ying-Yong Wang. Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China. Zoological Research, 2020, 41(2): 105-122. doi: 10.24272/j.issn.2095-8137.2020.033

短腿蟾属(无尾目:角蟾科)复查并恢复平头短腿蟾的有效性及描述一新种

doi: 10.24272/j.issn.2095-8137.2020.033

Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China

Funds: This work was supported by the Project of Comprehensive Scientific Survey of Luoxiao Mountains Region of Ministry of Science and Technology, China (2013FY111500); Specimen Platform of Ministry of Science and Technology, China, teaching specimens sub-platform (2005DKA21403-JK); Animal Branch of the Germplasm Bank of Wild Species, Chinese Academy of Sciences (Large Research Infrastructure Funding); and Project of Scientific Investigation on Amphibian, Reptilian and Avian Animals in Jiangxi Jiulianshan National Nature Reserve
More Information
  • 摘要:

    最近,关于广义角蟾属有诸多分类争论,其中也包括单系群短腿蟾。短腿蟾具有明显的可识别的形态特征,且最近所有分子系统学研究都表明其为一个稳定的单系群,因此将其作为一个独立属是合理的。本研究对该属做了全面复查,并基于形态学差异和系统发育关系提出了两个种组,同时认为平头短腿蟾是一个有效种,其与宽头短腿蟾有较大遗传分化和显著形态差异。此外,基于采自中国东南部的一系列标本,描述了一新种——东方短腿蟾。至此,短腿蟾属成员增加到7个,其多样性显然被低估了。另外,基于分类便利性、单系性和紧凑性原则,本研究还讨论了角蟾亚科其他单系群的属级分类地位问题。

  • Figure  1.  Collection localities of samples used in this study and habitat of Brachytarsophrys orientalis sp. nov.

    A: Localities of Brachytarsophrys orientalis sp. nov.: 1: Huboliao Nature Reserve, Fujian; 2: Shanghang County, Fujian; 3: Jiulianshan Nature Reserve, Jiangxi. Localities of B. popei; 4: Taoyuandong Nature Reserve, Hunan. Localities of B. chuannanensis; 5: Hejiang County, Sichuan. Localities of B. platyparietus; 6: Mt. Fanjing, Guizhou; 7: Mt. Jinzhong, Guangxi; 8: Shiping County, Yunnan; 9: Mt. Mopan, Yunnan; 10: Dayao County, Yunnan; 11: Yanbian County, Yunnan. Localities of B. feae; 12: Jingdong County, Yunnan. Localities of B. carinense; 13: Mae Surin NP., Mae Hong Son, Thailand; 14: Omkoi, Chiang Mai, Thailand; 15: Thong Pha Phum, Kanchanaburi, Thailand. Localities of B. intermedia; 16: Krong Pa, Gia Lai, Vietnam. B: Habitat of Brachytarsophrys orientalis sp. nov. in Jiulianshan Nature Reserve, Jiangxi Province.

    Figure  2.  Location of web on phalange articulation

    (4-) IV: Distal part of articulation between fourth phalange and metatarsal; (4) IV: Middle part of articulation between fourth phalange and metatarsal; (4+) IV: Proximal part of articulation between fourth phalange and metatarsal; (4++) IV: Lower part of articulation between fourth phalange and metatarsal.

    Figure  3.  Bayesian inference and maximum-likelihood phylogenies

    Numbers before slashes are Bayesian posterior probabilities, and numbers after slashes are maximum-likelihood bootstrap supports.

    Figure  4.  Boxplot of SVL showing body size variation among adult Brachytarsophrys males (data deficiency of B. chuannanensis and B. carinense)

    Horizontal lines within each box represent median, and boxes encompass 75th and 25th percentile.

    Figure  5.  General aspect of Brachytarsophrys chuannanensis

    A: Dorsolateral view of adult male holotype CIB 98A0045 in preservative; B: Ventral view of holotype CIB 98A0045 in preservative; C, D: Hand and foot of holotype CIB 98A0045 in preservative; E, F: Ventral view of 38th stage tadpole of B. chuannanensis.

    Figure  6.  General aspect of Brachytarsophrys feae

    A: Dorsolateral view of adult male SYS a003914 in life; B: Ventral view of adult male SYS a003914 in life; C, D: Hand and foot of adult male SYS a003914 in life; E: Ventral view of 44th stage tadpole of B. feae.

    Figure  7.  General aspect of Brachytarsophrys popei

    A: Dorsolateral view of adult male holotype SYS a001867 in life; B: Ventral view of holotype SYS a001867 in life.

    Figure  8.  General aspect of Brachytarsophrys platyparietus

    A: Dorsolateral view of adult male SYS a005919 in life; B: Ventral view of adult male SYS a005919 in life; C, D: Hand and foot of adult male SYS a005919 in life.

    Figure  9.  Numerous small, conical, horny tubercles in Brachytarsophrys platyparietus.

    A, B: Conical horny tubercles in adult male SYS a005919; C, D: Conical horny tubercles in adult female SYS a002236.

    Figure  10.  Egg protection behavior in Brachytarsophrys platyparietus

    A: Under rock, adult male on left protected eggs on right; B: Eggs of B. platyparietus.

    Figure  11.  General aspect of Brachytarsophrys orientalis sp. nov.

    A: Dorsolateral view of adult male holotype SYS a004227 in life; B: Ventral view of holotype SYS a004227 in life; C, D: Hand and foot of holotype SYS a004227 in life.

    Figure  12.  Morphological differences between Brachytarsophrys orientalis sp. nov. and B. popei

    A: Sole of feet in male holotype SYS a004227 of Brachytarsophrys orientalis sp. nov.; B: Sole of feet in female paratype SYS a004486 of Brachytarsophrys orientalis sp. nov.; C: Sole of feet in male holotype SYS a001867 of B. popei; D: Sole of feet in female paratype SYS a001875 of B. popei; E, F: Ventral view of 36th stage tadpole of Brachytarsophrys orientalis sp. nov.; G, H: Ventral view of 29th stage tadpole of B. popei.

    Table  1.   Localities, voucher information, and GenBank accession Nos. of all samples used in this study

    IDSpeciesLocalities (*: Type locality)Specimen No.COIcyt b
    1Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a004225MT162625MT162650
    2Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a004226MT162626MT162651
    3Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a004227MT162627MT162652
    4Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a004228MT162628MT162653
    5Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a004486MT162629MT162654
    6Brachytarsophrys orientalis sp. nov.*China: Jiulianshan Nature Reserve, Longnan County, JiangxiSYS a005451MT162632MT162655
    7Brachytarsophrys orientalis sp. nov.China: Gutian Township, Shanghang County, FujianSYS a003249MT162623MT162648
    8Brachytarsophrys orientalis sp. nov.China: Huboliao Nature Reserve, Nanjing County, FujianSYS a003340MT162624MT162649
    9Brachytarsophrys carinenseThailand: Doi Chiang Dao, Chiang MaiK3001KR087626
    10Brachytarsophrys carinenseThailand: Omkoi, Chiang MaiKIZ024170MT162640MT162663
    11Brachytarsophrys carinenseThailand: Mae Surin NP., Mae Hong SonKIZ024429MT162641MT162664
    12Brachytarsophrys carinenseThailand: Thong Pha Phum, KanchanaburiKIZ024640MT162642MT162665
    13Brachytarsophrys chuannanensis*China: Zihuai Township, Hejiang County, SichuanSYS a004926MT162630
    14Brachytarsophrys chuannanensis*China: Zihuai Township, Hejiang County, SichuanSYS a004927MT162631
    15Brachytarsophrys feaeChina: Jingdong County, YunnanSYS a003912MH406362MH407192
    16Brachytarsophrys feaeChina: Jingdong County, YunnanSYS a003913MH406363MH407193
    17Brachytarsophrys intermediaVietnam: Krong Pa, Gia LaiROM 23794MT162643MT162666
    18Brachytarsophrys platyparietus*China: Duodihe, Dayao county, YunnanSYS a005919MT162633MT162656
    19Brachytarsophrys platyparietusChina: Mt. Jinzhong, Longlin County, GuangxiSYS a002236MT162622MT162647
    20Brachytarsophrys platyparietusChina: Mt. Fanjing, Tongren City, GuizhouYPX43968MT162644MT162667
    21Brachytarsophrys platyparietusChina: Mt. Mopan, Xinping County, YunnanSYS a007774MT162634MT162657
    22Brachytarsophrys platyparietusChina: Mt. Mopan, Xinping County, YunnanSYS a007775MT162635MT162658
    23Brachytarsophrys platyparietusChina: Mt. Mopan, Xinping County, YunnanSYS a007776MT162636MT162659
    24Brachytarsophrys platyparietusChina: Mt. Mopan, Xinping County, YunnanSYS a007777MT162637MT162660
    25Brachytarsophrys platyparietusChina: Yilong Township, Shiping County, YunnanSYS a007790MT162638MT162661
    26Brachytarsophrys platyparietusChina: Yumen Township, Yanbian County, SichuanSYS a007853MT162639MT162662
    27Brachytarsophrys popei*China: Taoyuandong Nature Reserve, Yanling County, HunanSYS a001864MH406361MH407191
    28Brachytarsophrys popei*China: Taoyuandong Nature Reserve, Yanling County, HunanSYS a001865MT162620MT162645
    29Brachytarsophrys popei*China: Taoyuandong Nature Reserve, Yanling County, HunanSYS a001866MT162621MT162646
    30Atympanophrys shapingensisChina: Mt. Wawu, Hongya County, SichuanSYS a005310MH406352MH407182
    31Atympanophrys shapingensisChina: Zhaojue County, SichuanSYS a005339MH406359MH407189
    32Xenophrys mangshanensisChina: Mt. Dayao, Jinxiu County, GuangxiSYS a004870MH406323MH407153
    33Xenophrys mangshanensisChina: Mt. Dayao, Jinxiu County, GuangxiSYS a004871MH406324MH407154
    –: Not available.
    下载: 导出CSV

    Table  2.   Primer pairs used in this study

    GeneForward primerReverse primerReferences
    COIChmf4 (5'-TYTCWACWAAYCAYAAAGAYATCGG-3')Chmr4 (5'-ACYTCRGGRTGRCCRAARAATCA-3')Che et al., 2012
    COIDglco (5'-GGTCAACAAATCATAAAGAYATYGG-3')Dghco (5'-TAAACTTCAGGGTGACCAAARAAYCA-3')Meyer et al., 2005
    cyt bPFGlu14140L(5'-GAAAAACCACTGTTGTHHYTCAACTA-3')PFThr15310 (5'-CGGYTTACAAGACCGRTGCTTT-3')Zhang et al., 2013
    下载: 导出CSV

    Table  3.   Uncorrected P-distances (mean, in %) among species of the genus Brachytarsophrys based on partial mitochondrial COI gene

    IDSpecies1–89–1213–1415–161718–2627–29
    1–8Brachytarsophrys orientalis sp. nov.0.3
    9–12B. carinense15.80.6
    13–14B. chuannanensis8.514.70.5
    15–16B. feae9.415.36.30.0
    17B. intermedia15.712.116.315.2
    18–26B. platyparietus8.414.65.86.815.90.5
    27–29B. popei4.315.88.710.415.77.70.1
    下载: 导出CSV

    Table  4.   Measurements (in mm; minimum−maximum, mean±SD) and body proportions of examined specimens of Brachytarsophrys

    Brachytarsophrys orientalis sp. nov.B. popeiB. platyparietusB. chuannanensisB. feaeB. intermedia
    M (n=7)F (n=1)M (n=13)F (n=1)M (n=6)F (n=3)M (n=12)M (n=5)M (n=7)F (n=1)
    SVL76.8–82.7 (79.9±2.6)88.670.7–83.5 (76.9±3.5)86.288.5–113.0 (101.8±9.6)118.5–131.0 (124.8±6.3)91.4–109.4 (105.3 in CIB 98A0045)78.5–94.9 (86.9±7.4)86.0–103.0 (95.6±6.1)92.0
    HDL32.8–35.0 (33.6±0.7)40.829.8–34.2 (32.2±1.3)36.037.1–47.0 (42.0±3.9)46.5–52.5 (49.1±3.1)44.8 in CIB 98A004531.5–35.9 (33.8±1.6)
    SKL20.1–21.5 (20.9±0.5)24.617.7–20.6 (19.6±1.0)21.223.9 in SYS a00591930.6 in SYS a00223621.0–23.3 (22.1±0.9)23.0–28.0 (25.3±1.5)27.0
    HDW39.1–42.3 (40.8±1.4)48.836.0–40.8 (39.1±1.6)42.543.1–57.0 (50.8±5.2)55.7–61.0 (58.9±2.8)54.1 in CIB 98A004537.7–44.4 (41.3±2.4)44.0–51.0 (47.3±2.3)50.0
    SNT9.0–10.6 (9.8±0.6)12.28.5–10.3 (9.6±0.5)10.69.9 in SYS a00591913.5 in SYS a0022369.2–10.5 (9.7±0.5)11.0–13.0 (12.0±0.6)12.0
    IND7.6–9.0 (8.4±0.5)9.57.0–8.7 (7.8±0.4)8.28.2 in SYS a00591910.3 in SYS a0022367.3–8.7 (8.3±0.6)
    IOD8.9–13.6 (12.0±1.7)14.611.1–14.0 (12.0±1.2)13.910.6 in SYS a00591914.0 in SYS a0022368.8–13.9 (11.0±2.0)12.0–13.0 (12.4±0.5)13.0
    ED8.5–10.7 (9.8±0.9)10.47.8–10.6 (9.3±1.0)9.88.7 in SYS a0059199.9 in SYS a0022367.6–11.3 (9.1±1.4)
    HND20.0–22.5 (21.3±0.8)22.618.6–21.6 (19.8±1.0)20.526.7 in SYS a00591932.1 in SYS a00223621.3–24.0 (22.7±1.0)23.0–27.0 (24.4±1.3)25.0
    RAD14.7–19.7 (17.5±1.7)17.415.6–17.9 (16.9±0.8)17.317.6 in SYS a00591923.2 in SYS a00223617.0–19.1 (17.8±0.9)
    TIB31.1–33.5 (32.1±1.0)33.128.0–33.5 (30.8±1.6)31.434.3 in SYS a00591943.8 in SYS a00223642.7 in CIB 98A004531.3–36.6 (33.6±1.9)32.0–39.0 (35.9±2.1)36.0
    FTL46.9–50.5 (48.4±1.1)51.042.0–49.7 (46.1±2.3)49.557.4 in SYS a00591970.6 in SYS a00223647.1–50.9 (48.9±1.4)
    HDL/SVL0.41–0.43 (0.42±0.01)0.460.40–0.43 (0.42±0.01)0.420.39–0.45 (0.41±0.02)0.39–0.40 (0.39±0.01)0.43 in CIB 98A00450.37–0.42 (0.39±0.02)
    HDW/SVL0.49–0.53 (0.51±0.01)0.550.47–0.53 (0.51±0.02)0.490.46–0.54 (0.50±0.03)0.47–0.48 (0.47±0.01)0.51 in CIB 98A00450.45–0.53 (0.48±0.03)0.47–0.51 (0.50±0.02)0.54
    HDW/SKL1.89–2.02 (1.95±0.05)1.981.89–2.24 (2.00±0.09)2.001.80 in SYS a0059191.96 in SYS a0022361.73–1.96 (1.87±0.10)1.82–1.92 (1.87±0.04)1.85
    HDW/HDL1.17–1.26 (1.22±0.04)1.201.18–1.26 (1.21±0.03)1.181.16–1.24 (1.21±0.03)1.16–1.24 (1.20±0.04)1.21 in CIB 98A00451.20–1.25 (1.22±0.02)
    SNT/HDL0.27–0.31 (0.29±0.01)0.300.28–0.31 (0.30±0.01)0.290.27 in SYS a0059190.28 in SYS a0022360.27–0.31 (0.29±0.02)
    SNT/SVL0.12–0.13 (0.12±0.01)0.140.12–0.13 (0.13±0.00)0.120.10 in SYS a0059190.11 in SYS a0022360.10–0.12 (0.11±0.01)0.12–0.13 (0.13±0.00)0.13
    IND/IOD0.60–0.92 (0.71±0.11)0.650.60–0.71 (0.65±0.03)0.590.77 in SYS a0059190.74 in SYS a0022360.61–0.94 (0.77±0.15)
    IND/HDW0.19–0.23 (0.21±0.02)0.190.18–0.21 (0.20±0.01)0.190.19 in SYS a0059190.17 in SYS a0022360.18–0.22 (0.20±0.01)
    IOD/HDW0.22–0.32 (0.29±0.04)0.300.29–0.35 (0.31±0.02)0.330.25 in SYS a0059190.23 in SYS a0022360.23–0.31 (0.26±0.03)
    ED/HDL0.26–0.32 (0.29±0.02)0.250.25–0.33 (0.29±0.02)0.270.23 in SYS a0059190.20 in SYS a0022360.24–0.31 (0.27±0.03)
    ED/SVL0.11–0.14 (0.12±0.01)0.120.11–0.14 (0.12±0.01)0.110.09 in SYS a0059190.08 in SYS a0022360.09–0.12 (0.10±0.02)
    HND/SVL0.25–0.28 (0.27±0.01)0.260.24–0.28 (0.26±0.01)0.240.28 in SYS a0059190.26 in SYS a0022360.25–0.29 (0.26±0.02)0.23–0.27 (0.26±0.01)0.27
    RAD/SVL0.19–0.24 (0.22±0.02)0.200.20–0.24 (0.22±0.01)0.200.19 in SYS a0059190.19 in SYS a0022360.19–0.23 (0.21±0.02)
    TIB/SVL0.39–0.42 (0.40±0.01)0.370.38–0.43 (0.40±0.01)0.360.36 in SYS a0059190.35 in SYS a0022360.41 in CIB 98A00450.36–0.42 (0.39±0.02)0.35–0.41 (0.37±0.02)0.39
    FTL/SVL0.58–0.65 (0.61±0.03)0.580.57–0.63 (0.60±0.02)0.570.61 in SYS a0059190.57 in SYS a0022360.53–0.61 (0.57±0.03)
    ReferencesThis studyZhao et al., 2014Rao & Yang, 1997;
    this study
    Fei & Ye, 2001; this studyThis studySmith, 1921
    F: Female; M: Male. –: Not available.
    下载: 导出CSV

    Table  5.   Diagnostic characters separating the seven Brachytarsophrys species from each other

    Morphological charactersBrachytarsophrys orientalis sp. nov.B. popeiB. carinenseB. platyparietusB. chuannanensisB. feaeB. intermedia
    SVL of males (in mm)76.8–82.770.7–83.591.6–123.088.5–113.091.4–109.478.5–94.986.0–103.0
    SVL of females (in mm)88.686.2124.0–168.0118.5–131.092.0
    TongueFeebly notched behindDeeply notched behindFeebly notched behindFeebly notched behindFeebly notched behindFeebly notched behindFeebly notched behind
    Dermal ridge or glandular fold on dorsumAbsentAbsentDermal ridge on dorsumAbsentAbsentAbsentGlandular fold on dorsum
    Small, conical, horny tuberclesAbsentAbsentAbsentPresent on pectoral region, lateral belly to lower flank of body, ventral surface, and rear of limbsAbsentAbsentAbsent
    Stellate bonyAbsentAbsentStellate bony deposits in skin of parietal region and anterior part of dorsumAbsentAbsentStellate bony deposits on each side of parietal regionAbsent
    Webbing formula of maleI (1½)-(2) II (1½)-(3) III (2½)-(4) IV (4)-(2) VI (1½)-(2) II (1½)-(3) III (2½)-(3⅔) IV (3⅔)-(2) VI (1½)-(2+) II (1½)-(3) III (2⅓)-(3⅔) IV (3⅔)-(2-) VI (1½)-(2++) II (2)-(3++) III (3)-(4) IV (4++)-(2½) VI (2)-(2++) II (2-)-(3) III (2⅔)-(4) IV (4)-(2⅔) V
    Webbing formula of femaleI (2)-(2+) II (1⅔)-(3) III (3-)-(4) IV (4+)-(2½) VI (1½)-(2+) II (1½)-(3) III (2½)-(4-) IV (4-)-(2) VI (1½)-(2+) II (1½)-(3) III (2⅔)-(4-) IV (4-)-(2⅔) V
    Lateral fringes on toes of maleOne third as broad as distal toe phalanxOne fourth as broad as distal toe phalanxHalf as broad as distal toe phalanxOne fifth as broad as distal toe phalanx in CIB 98A0045One fourth as broad as distal toe phalanx
    Lateral fringes on toes of femaleOne sixth as broad as distal toe phalanxOne fifth as broad as distal toe phalanxOne fourth as broad as distal toe phalanx
    Position of tibiotarsal articulation reachingCommissure of jawCommissure of jawAxilla in females, angle of mouth in malesCommissure of jawShoulder or commissure of jawShoulder in females, commissure of jaw in malesCommissure of jaw
    Transversal stripe on chest in tadpoleAbsentA transversal stripe on chestA transversal stripe on chestSeveral transversal stripes on chest
    –: Not available.
    下载: 导出CSV
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