Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China
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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。
Abstract:The genus-level recognition of monophyletic short-legged toads (Brachytarsophrys) has been recently implicated in the taxonomic debate of Megophrys sensu lato. In the present study, Brachytarsophrys is reasonably regarded as a distinct genus based on significant morphological differentiations and recent molecular analyses. Furthermore, a comprehensive review of this genus is performed, with two species groups proposed based on morphological differences and phylogenetic relationships. Particularly, Brachytarsophrys platyparietus is removed as a synonym of Brachytarsophrys carinense and considered a valid species due to significant genetic divergence and distinct morphological differences. In addition, a new species, Brachytarsophrys orientalis sp . nov ., is described based on a series of specimens collected from southeastern China. This work takes the member species of the genus Brachytarsophrys to seven, suggesting that the diversity of Brachytarsophrys is underestimated. In addition, the genus levels of other monophyletic groups within the subfamily Megophryinae are discussed.
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Key words:
- Genus level /
- Megophryinae /
- Morphology /
- Phylogeny /
- Revision
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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
ID Species Localities (*: Type locality) Specimen No. COI cyt b 1 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a004225 MT162625 MT162650 2 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a004226 MT162626 MT162651 3 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a004227 MT162627 MT162652 4 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a004228 MT162628 MT162653 5 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a004486 MT162629 MT162654 6 Brachytarsophrys orientalis sp. nov. *China: Jiulianshan Nature Reserve, Longnan County, Jiangxi SYS a005451 MT162632 MT162655 7 Brachytarsophrys orientalis sp. nov. China: Gutian Township, Shanghang County, Fujian SYS a003249 MT162623 MT162648 8 Brachytarsophrys orientalis sp. nov. China: Huboliao Nature Reserve, Nanjing County, Fujian SYS a003340 MT162624 MT162649 9 Brachytarsophrys carinense Thailand: Doi Chiang Dao, Chiang Mai K3001 KR087626 – 10 Brachytarsophrys carinense Thailand: Omkoi, Chiang Mai KIZ024170 MT162640 MT162663 11 Brachytarsophrys carinense Thailand: Mae Surin NP., Mae Hong Son KIZ024429 MT162641 MT162664 12 Brachytarsophrys carinense Thailand: Thong Pha Phum, Kanchanaburi KIZ024640 MT162642 MT162665 13 Brachytarsophrys chuannanensis *China: Zihuai Township, Hejiang County, Sichuan SYS a004926 MT162630 – 14 Brachytarsophrys chuannanensis *China: Zihuai Township, Hejiang County, Sichuan SYS a004927 MT162631 – 15 Brachytarsophrys feae China: Jingdong County, Yunnan SYS a003912 MH406362 MH407192 16 Brachytarsophrys feae China: Jingdong County, Yunnan SYS a003913 MH406363 MH407193 17 Brachytarsophrys intermedia Vietnam: Krong Pa, Gia Lai ROM 23794 MT162643 MT162666 18 Brachytarsophrys platyparietus *China: Duodihe, Dayao county, Yunnan SYS a005919 MT162633 MT162656 19 Brachytarsophrys platyparietus China: Mt. Jinzhong, Longlin County, Guangxi SYS a002236 MT162622 MT162647 20 Brachytarsophrys platyparietus China: Mt. Fanjing, Tongren City, Guizhou YPX43968 MT162644 MT162667 21 Brachytarsophrys platyparietus China: Mt. Mopan, Xinping County, Yunnan SYS a007774 MT162634 MT162657 22 Brachytarsophrys platyparietus China: Mt. Mopan, Xinping County, Yunnan SYS a007775 MT162635 MT162658 23 Brachytarsophrys platyparietus China: Mt. Mopan, Xinping County, Yunnan SYS a007776 MT162636 MT162659 24 Brachytarsophrys platyparietus China: Mt. Mopan, Xinping County, Yunnan SYS a007777 MT162637 MT162660 25 Brachytarsophrys platyparietus China: Yilong Township, Shiping County, Yunnan SYS a007790 MT162638 MT162661 26 Brachytarsophrys platyparietus China: Yumen Township, Yanbian County, Sichuan SYS a007853 MT162639 MT162662 27 Brachytarsophrys popei *China: Taoyuandong Nature Reserve, Yanling County, Hunan SYS a001864 MH406361 MH407191 28 Brachytarsophrys popei *China: Taoyuandong Nature Reserve, Yanling County, Hunan SYS a001865 MT162620 MT162645 29 Brachytarsophrys popei *China: Taoyuandong Nature Reserve, Yanling County, Hunan SYS a001866 MT162621 MT162646 30 Atympanophrys shapingensis China: Mt. Wawu, Hongya County, Sichuan SYS a005310 MH406352 MH407182 31 Atympanophrys shapingensis China: Zhaojue County, Sichuan SYS a005339 MH406359 MH407189 32 Xenophrys mangshanensis China: Mt. Dayao, Jinxiu County, Guangxi SYS a004870 MH406323 MH407153 33 Xenophrys mangshanensis China: Mt. Dayao, Jinxiu County, Guangxi SYS a004871 MH406324 MH407154 –: Not available. 
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Table 2. Primer pairs used in this study
Gene Forward primer Reverse primer References COI Chmf4 (5'-TYTCWACWAAYCAYAAAGAYATCGG-3') Chmr4 (5'-ACYTCRGGRTGRCCRAARAATCA-3') Che et al., 2012 COI Dglco (5'-GGTCAACAAATCATAAAGAYATYGG-3') Dghco (5'-TAAACTTCAGGGTGACCAAARAAYCA-3') Meyer et al., 2005 cyt b PFGlu14140L(5'-GAAAAACCACTGTTGTHHYTCAACTA-3') PFThr15310 (5'-CGGYTTACAAGACCGRTGCTTT-3') Zhang et al., 2013
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Table 3. Uncorrected P-distances (mean, in %) among species of the genus Brachytarsophrys based on partial mitochondrial COI gene
ID Species 1–8 9–12 13–14 15–16 17 18–26 27–29 1–8 Brachytarsophrys orientalis sp. nov. 0.3 9–12 B. carinense 15.8 0.6 13–14 B. chuannanensis 8.5 14.7 0.5 15–16 B. feae 9.4 15.3 6.3 0.0 17 B. intermedia 15.7 12.1 16.3 15.2 – 18–26 B. platyparietus 8.4 14.6 5.8 6.8 15.9 0.5 27–29 B. popei 4.3 15.8 8.7 10.4 15.7 7.7 0.1
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Table 4. Measurements (in mm; minimum−maximum, mean±SD) and body proportions of examined specimens of Brachytarsophrys
Brachytarsophrys orientalis sp. nov. B. popei B. platyparietus B. chuannanensis B. feae B. 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) SVL 76.8–82.7 (79.9±2.6) 88.6 70.7–83.5 (76.9±3.5) 86.2 88.5–113.0 (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 HDL 32.8–35.0 (33.6±0.7) 40.8 29.8–34.2 (32.2±1.3) 36.0 37.1–47.0 (42.0±3.9) 46.5–52.5 (49.1±3.1) 44.8 in CIB 98A0045 31.5–35.9 (33.8±1.6) – – SKL 20.1–21.5 (20.9±0.5) 24.6 17.7–20.6 (19.6±1.0) 21.2 23.9 in SYS a005919 30.6 in SYS a002236 – 21.0–23.3 (22.1±0.9) 23.0–28.0 (25.3±1.5) 27.0 HDW 39.1–42.3 (40.8±1.4) 48.8 36.0–40.8 (39.1±1.6) 42.5 43.1–57.0 (50.8±5.2) 55.7–61.0 (58.9±2.8) 54.1 in CIB 98A0045 37.7–44.4 (41.3±2.4) 44.0–51.0 (47.3±2.3) 50.0 SNT 9.0–10.6 (9.8±0.6) 12.2 8.5–10.3 (9.6±0.5) 10.6 9.9 in SYS a005919 13.5 in SYS a002236 – 9.2–10.5 (9.7±0.5) 11.0–13.0 (12.0±0.6) 12.0 IND 7.6–9.0 (8.4±0.5) 9.5 7.0–8.7 (7.8±0.4) 8.2 8.2 in SYS a005919 10.3 in SYS a002236 – 7.3–8.7 (8.3±0.6) – – IOD 8.9–13.6 (12.0±1.7) 14.6 11.1–14.0 (12.0±1.2) 13.9 10.6 in SYS a005919 14.0 in SYS a002236 – 8.8–13.9 (11.0±2.0) 12.0–13.0 (12.4±0.5) 13.0 ED 8.5–10.7 (9.8±0.9) 10.4 7.8–10.6 (9.3±1.0) 9.8 8.7 in SYS a005919 9.9 in SYS a002236 – 7.6–11.3 (9.1±1.4) – – HND 20.0–22.5 (21.3±0.8) 22.6 18.6–21.6 (19.8±1.0) 20.5 26.7 in SYS a005919 32.1 in SYS a002236 – 21.3–24.0 (22.7±1.0) 23.0–27.0 (24.4±1.3) 25.0 RAD 14.7–19.7 (17.5±1.7) 17.4 15.6–17.9 (16.9±0.8) 17.3 17.6 in SYS a005919 23.2 in SYS a002236 – 17.0–19.1 (17.8±0.9) – – TIB 31.1–33.5 (32.1±1.0) 33.1 28.0–33.5 (30.8±1.6) 31.4 34.3 in SYS a005919 43.8 in SYS a002236 42.7 in CIB 98A0045 31.3–36.6 (33.6±1.9) 32.0–39.0 (35.9±2.1) 36.0 FTL 46.9–50.5 (48.4±1.1) 51.0 42.0–49.7 (46.1±2.3) 49.5 57.4 in SYS a005919 70.6 in SYS a002236 – 47.1–50.9 (48.9±1.4) – – HDL/SVL 0.41–0.43 (0.42±0.01) 0.46 0.40–0.43 (0.42±0.01) 0.42 0.39–0.45 (0.41±0.02) 0.39–0.40 (0.39±0.01) 0.43 in CIB 98A0045 0.37–0.42 (0.39±0.02) – – HDW/SVL 0.49–0.53 (0.51±0.01) 0.55 0.47–0.53 (0.51±0.02) 0.49 0.46–0.54 (0.50±0.03) 0.47–0.48 (0.47±0.01) 0.51 in CIB 98A0045 0.45–0.53 (0.48±0.03) 0.47–0.51 (0.50±0.02) 0.54 HDW/SKL 1.89–2.02 (1.95±0.05) 1.98 1.89–2.24 (2.00±0.09) 2.00 1.80 in SYS a005919 1.96 in SYS a002236 – 1.73–1.96 (1.87±0.10) 1.82–1.92 (1.87±0.04) 1.85 HDW/HDL 1.17–1.26 (1.22±0.04) 1.20 1.18–1.26 (1.21±0.03) 1.18 1.16–1.24 (1.21±0.03) 1.16–1.24 (1.20±0.04) 1.21 in CIB 98A0045 1.20–1.25 (1.22±0.02) – – SNT/HDL 0.27–0.31 (0.29±0.01) 0.30 0.28–0.31 (0.30±0.01) 0.29 0.27 in SYS a005919 0.28 in SYS a002236 – 0.27–0.31 (0.29±0.02) – – SNT/SVL 0.12–0.13 (0.12±0.01) 0.14 0.12–0.13 (0.13±0.00) 0.12 0.10 in SYS a005919 0.11 in SYS a002236 – 0.10–0.12 (0.11±0.01) 0.12–0.13 (0.13±0.00) 0.13 IND/IOD 0.60–0.92 (0.71±0.11) 0.65 0.60–0.71 (0.65±0.03) 0.59 0.77 in SYS a005919 0.74 in SYS a002236 0.61–0.94 (0.77±0.15) – – IND/HDW 0.19–0.23 (0.21±0.02) 0.19 0.18–0.21 (0.20±0.01) 0.19 0.19 in SYS a005919 0.17 in SYS a002236 – 0.18–0.22 (0.20±0.01) – – IOD/HDW 0.22–0.32 (0.29±0.04) 0.30 0.29–0.35 (0.31±0.02) 0.33 0.25 in SYS a005919 0.23 in SYS a002236 – 0.23–0.31 (0.26±0.03) – – ED/HDL 0.26–0.32 (0.29±0.02) 0.25 0.25–0.33 (0.29±0.02) 0.27 0.23 in SYS a005919 0.20 in SYS a002236 – 0.24–0.31 (0.27±0.03) – – ED/SVL 0.11–0.14 (0.12±0.01) 0.12 0.11–0.14 (0.12±0.01) 0.11 0.09 in SYS a005919 0.08 in SYS a002236 – 0.09–0.12 (0.10±0.02) – – HND/SVL 0.25–0.28 (0.27±0.01) 0.26 0.24–0.28 (0.26±0.01) 0.24 0.28 in SYS a005919 0.26 in SYS a002236 – 0.25–0.29 (0.26±0.02) 0.23–0.27 (0.26±0.01) 0.27 RAD/SVL 0.19–0.24 (0.22±0.02) 0.20 0.20–0.24 (0.22±0.01) 0.20 0.19 in SYS a005919 0.19 in SYS a002236 – 0.19–0.23 (0.21±0.02) – – TIB/SVL 0.39–0.42 (0.40±0.01) 0.37 0.38–0.43 (0.40±0.01) 0.36 0.36 in SYS a005919 0.35 in SYS a002236 0.41 in CIB 98A0045 0.36–0.42 (0.39±0.02) 0.35–0.41 (0.37±0.02) 0.39 FTL/SVL 0.58–0.65 (0.61±0.03) 0.58 0.57–0.63 (0.60±0.02) 0.57 0.61 in SYS a005919 0.57 in SYS a002236 – 0.53–0.61 (0.57±0.03) – – References This study Zhao et al., 2014 Rao & Yang, 1997;
this studyFei & Ye, 2001; this study This study Smith, 1921 F: Female; M: Male. –: Not available.
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Table 5. Diagnostic characters separating the seven Brachytarsophrys species from each other
Morphological characters Brachytarsophrys orientalis sp. nov. B. popei B. carinense B. platyparietus B. chuannanensis B. feae B. intermedia SVL of males (in mm) 76.8–82.7 70.7–83.5 91.6–123.0 88.5–113.0 91.4–109.4 78.5–94.9 86.0–103.0 SVL of females (in mm) 88.6 86.2 124.0–168.0 118.5–131.0 – – 92.0 Tongue Feebly notched behind Deeply notched behind Feebly notched behind Feebly notched behind Feebly notched behind Feebly notched behind Feebly notched behind Dermal ridge or glandular fold on dorsum Absent Absent Dermal ridge on dorsum Absent Absent Absent Glandular fold on dorsum Small, conical, horny tubercles Absent Absent Absent Present on pectoral region, lateral belly to lower flank of body, ventral surface, and rear of limbs Absent Absent Absent Stellate bony Absent Absent Stellate bony deposits in skin of parietal region and anterior part of dorsum Absent Absent Stellate bony deposits on each side of parietal region Absent Webbing formula of male I (1½)-(2) II (1½)-(3) III (2½)-(4) IV (4)-(2) V I (1½)-(2) II (1½)-(3) III (2½)-(3⅔) IV (3⅔)-(2) V – I (1½)-(2+) II (1½)-(3) III (2⅓)-(3⅔) IV (3⅔)-(2-) V I (1½)-(2++) II (2)-(3++) III (3)-(4) IV (4++)-(2½) V I (2)-(2++) II (2-)-(3) III (2⅔)-(4) IV (4)-(2⅔) V – Webbing formula of female I (2)-(2+) II (1⅔)-(3) III (3-)-(4) IV (4+)-(2½) V I (1½)-(2+) II (1½)-(3) III (2½)-(4-) IV (4-)-(2) V – I (1½)-(2+) II (1½)-(3) III (2⅔)-(4-) IV (4-)-(2⅔) V – – – Lateral fringes on toes of male One third as broad as distal toe phalanx One fourth as broad as distal toe phalanx – Half as broad as distal toe phalanx One fifth as broad as distal toe phalanx in CIB 98A0045 One fourth as broad as distal toe phalanx – Lateral fringes on toes of female One sixth as broad as distal toe phalanx One fifth as broad as distal toe phalanx – One fourth as broad as distal toe phalanx – – – Position of tibiotarsal articulation reaching Commissure of jaw Commissure of jaw Axilla in females, angle of mouth in males Commissure of jaw Shoulder or commissure of jaw Shoulder in females, commissure of jaw in males Commissure of jaw Transversal stripe on chest in tadpole Absent A transversal stripe on chest – – A transversal stripe on chest Several transversal stripes on chest – –: Not available.
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