Zoosyst. Evol. 100 (3) 2024, 1121-1134 | DOI 10.3897/zse.100.127268 > PENSUFT. Gy MuseuM ror BERLIN Taxonomic determination of Hypselotriton populations distributed in eastern Guangdong, China (Caudata, Salamandridae), with description of a new species and a new subgenus Jian Wang! *, Zhao-Chi Zeng! , Tian-Li Wei!, Zhi-Tong Lyu* 1 Guangdong Polytechnic of Environmental Protection Engineering, Foshan 528216, Guangdong, China 2 The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, Guangdong, China 3 CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610040, Sichuan, China https://zoobank. org/7720F592-80B8-417D-AB7F-EF 28AE0B91B4 Corresponding authors: Jian Wang (wangj-1994@outlook.com); Zhi-Tong Lyu (lvzt@foxmail.com) Academic editor: Umilaela Arifin # Received 10 May 2024 Accepted 22 July 2024 # Published 14 August 2024 Abstract In this work, the Hypselotriton populations distributed in eastern Guangdong, China are studied in detail to clarify their taxonom- ic status. Based on morphological comparison and phylogenetic analysis, H. glaucus syn. nov. is synonymised with H. orphicus. Hypselotriton (Cynotriton) oolong sp. nov. from Mt Fenghuang in Chaozhou which used to be misidentified as H. orphicus, is re- vealed to be an independent lineage of subgenus Cynotriton and can be distinguished from all known congeners in morphology. By contrast, H. orphicus did not cluster within Cynotriton, but gathered with H. jiaoren comb. nov. to form a distinct unnamed clade within the genus. We therefore re-delimitate the intrageneric classification of the genus and a new subgenus Hakkatriton subgen. nov. is erected, corresponding to this unnamed clade. The Chinese Fire-bellied Newt genus Hypselotriton currently contains three subgen- era and about ten known species. Identified keys to the subgenera and related congeners of genus Hypse/otriton are further provided. Key Words Chaozhou, Cynops, Hakkatriton subgen. nov., Hypselotriton (Cynotriton) oolong sp. nov., Jiexi Introduction The Fire-bellied Newts contain about 18 species distrib- uted in China and Japan in East Asia (Raffaélli 2022; Lyu et al. 2023a). In traditional configuration, these spe- cies were provisionally placed in a single genus Cynops Tschudi, 1838, in spite of several controversies (Zhao and Hu 1984; Zhao et al. 1988; Lyu et al. 2023a). How- ever, recent phylogenetic studies suggested the Chinese and Japanese congeners are paraphyletic from each other (Rancilhac et al. 2021; Yuan et al. 2022). In the phyloge- netic analysis, the Fire-bellied Newt species distributed in Japanese Archipelago are revealed to be the basal lineage of the Modern Asian Newts. Compared with the insular species, the Fire-bellied Newt species occurring in main- land China are phylogenetically closer to other newt gen- era from mainland China and Indochina, 1.e. Pachytriton Boulenger, 1878, Paramesotriton Chang, 1935 and Lao- triton Dubois & Raffaélli, 2009 (Rancilhac et al. 2021; Yuan et al. 2022). Thus, Raffaélli (2022) partitioned these Species into two independent genera, Cynops for the Jap- anese Fire-bellied Newts and Hypselotriton Wolterstorff, 1934 for the Chinese Fire-bellied Newts and which was followed in this work. Particularly, a recent work has de- scribed two new species of Chinese Fire-bellied Newts (Lyu et al. 2023a), but did not adopt the latest taxonomic proposal by Raffaélli (2022) in a timely manner, result- ing in two new nomenclature combinations in this work, H. jiaoren (Lyu, Qi & Wang, 2023), comb. nov. and H. maguae (Lyu, Qi & Wang, 2023), comb. nov. Copyright Wang, J. et al. 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. 12 Within the genus Hypselotriton, Dubois and Raffaél- li (2011) classified the congeners into two subgenera, based on morphological and geographical characteristics, Hypselotriton and Cynotriton Dubois & Raffaélli, 2011, corresponding to the former Cynops wolterstorffi and Cynops orientalis groups, respectively (Zhao et al. 1988; Raffaélli 2022). Nonetheless, phylogenetic analysis has suggested three distinct and divergent clades within the genus, indicating that the intrageneric classification of this genus requires re-delimitation (Lyu et al. 2023a). The species diversity of Fire-bellied Newts was con- sidered underestimated. Tominaga et al. (2013, 2015) have revealed multiple distinct lineages within Cynops pyrrhogaster (Boie, 1826). Compared with the traditional recognition of two known species for Japanese Fire-bellied Newts, Raffaélli (2022) documented four nominated spe- cies and four unnamed species within the insular genus Cy- nops. Meanwhile, four new species of Hypselotriton have been described from Southeast Chinese Hilly Area since 2010, dramatically raising the diversity of this mainland genus (Wu et al. 2010; Yuan et al. 2013; Lyu et al. 2023a). However, the taxonomic status for several Hypselotriton Species remains unresolved. Lyu et al. (2023a) have dis- cussed the taxonomic confusion on the congeners from Yunnan-Guizhou Plateau in southwestern China that still require further studies. During our fieldwork and study on the Hypselotriton populations from Guangdong in south- eastern China, we have found that the recognition on H. or- Dhicus (Risch, 1983) is also with confusion, especially for its delimitation from another congener tn eastern Guang- dong, H. glaucus (Yuan, Jiang, Ding, Zhang & Che, 2013). Hypselotriton orphicus was nominated by Risch (1983), based on specimens collected and primarily described by Gressitt (1941) from “Dayang (Tai- Yong), Shantou Region [now belonging to Jiexi County, Jieyang City], 23°35'N, LIS°SI'E [= 23.58°N, 115.85°EF], altitude 640 m” (Fig. 1, sites 2-3). Hypselotriton glaucus was described, based on specimens collected from “Meiguang Village (23.67°N, 115.80°E; elevation 742 m), in Mt Lianhua, Wuhua Coun- ty, Meizhou” (Fig. 1, site 9) (Yuan et al. 2013), where it is in close proximity to the type locality of H. orphicus. When proposing H. glaucus, the data from Jiexi County were not mentioned. Instead, two separate populations col- lected from Mt Fenghuang of Guangdong and Mt Daiyun of Fujian were recorded as H. orphicus and used for com- parison (Yuan et al. 2013). Morphologically, the diagnostic characters of H. glaucus almost match the description of H. orphicus in Risch (1983), except for the irregular grey- ish-blue patches on the dorsum of H. glaucus. Such colour pattern was not described by Gressitt (1941) based on liv- ing or freshly-preserved specimens and would fade after preservation. Particularly, Risch (1983) has mentioned that the type series of H. orphicus are morphologically differ- ent from the population from central Fujian. Thus, the em- ployment of specimens from Mt Daryun, central Fujian (as well as those from Mt Fenghuang, eastern Guangdong) as H. orphicus might be problematic, which further adds to the confusion on the proposal of H. glaucus. zse.pensoft.net Wang, J. et al.: A new species and a new subgenus of Hypselotriton In this work, we perform morphological comparisons and molecular analyses on the topotypical population of Hypselotriton orphicus and H. glaucus, as well as on the Hypselotriton population from Mt Fenghuang (Fig. 1, site 1). The results suggest that H. glaucus is conspecific with H. orphicus, while the population from Mt Fenghuang represents an unnamed lineage of genus Hypselotriton that is described hereby. The taxonomic status for the population from central Fujian is also discussed. Further- more, we re-delimitate the intrageneric classification of the genus, as well as proposing a new subgenus for the clade comprised of H. orphicus and H. jiaoren comb. nov. Materials and methods Specimens and morphological analyses A series of museum specimens of the genus Hypselotriton from eastern Guangdong were examined. Detailed infor- mation for these specimens is presented in related species accounts below. Abbreviations for museums and institutes include: GEP (Guangdong Polytechnic of Environmental Protection Engineering, Foshan, China), CIB (Herpeto- logical Museum, Chengdu Institute of Biology, the Chi- nese Academy of Sciences, Chengdu, China), SYS (The Museum of Biology, Sun Yat-sen University, Guangzhou, China), MVZ (Museum of Vertebrate Zoology, University of California, Berkeley, USA), MNHN (Museum nation- al d’Histoire naturelle, Paris, France), AMNH (American Museum of Natural History, New York, USA) and CAS (California Academy of Sciences, San Francisco, USA). External measurements were made for the unnamed specimens with digital calipers (Neiko 01407A Stainless Steel 6-Inch Digital Caliper) to the nearest 0.1 mm. These measurements are as follows: total length (TOL) from tip of snout to tip of tail; snout—vent length (SVL) from tip of snout to posterior edge of vent; tail length (TAL) from posterior edge of vent to tip of tail; maximum tail depth (TAD); head length (HL) from tip of snout to the pos- terior edge of the parotoid gland; maximum head width (HW); snout length (SL) from tip of snout to the anterior corner of eye; eye diameter (ED) from the anterior cor- ner to the posterior corner of the eye; interorbital distance (IOD) between the anterior corner of each eye; eye—nos- tril length (EN) from the anterior corner of the eye to the nostril; internasal distance (IND) between the external nares; axilla—groin length (AG) between the axilla and the groin along the body; fore-limb length (FLL) from elbow to tip of finger III; and hind-limb length (HLL) from knee to tip of toe III. The morphological comparisons for known Hypselo- triton congeners were attained from literature of the orig- inal and subsequently supplemental descriptions (David 1873; Boulenger 1905; Gressitt 1941; Liu et al. 1962; Kou and Xing 1983; Risch 1983; Yang 1983; Fei and Ye 1983, 2016; Fei et al. 2006; Wu et al. 2010; Yuan et al. 2013; Raffaélli 2022; Lyu et al. 2023a). Zoosyst. Evol. 100 (3) 2024, 1121-1134 4 Hakkatriton subgen. nov. @ orphicus Cynotriton oolong sp. nov. » sp. glaucus syn. nov. ® @ fudingensis jiaoren comb. nov. @ maguae comb. nov. © orientalis Lis Jiangsu ys Shanghai Plot ae Jiangxi as t v Fujian Guangdong é fHong Kong Macao ey Hainan 100 km 115 120 Figure 1. Map showing the localities for congeners of genus Hypselotriton from Southeast Chinese Hilly Area. Inset map on top left zooms in eastern Guangdong Province, showing the localities for H. (Cynotriton) oolong sp. nov. and H. (Hakkatriton) orphicus: 1. Mt Feng- huang, Chao’an District, Chaozhou City (type locality of H. oolong sp. nov.); 2. Dayang Township, Jiexi County, Jieyang City (purported type locality of H. orphicus, see Remarks in the species account for locality delimitation); 3. Liangtian Township, Jiexi County (type local- ity of H. orphicus according to the original coordinate); 4. Mt Dabei, Jiexi County; 5. Mt Liwangzhang, Jiexi County; 6. Mt Hongtuzhang, Fengshun County, Meizhou City; 7. Mt Tongguzhang, Fengshun County; 8. Mt Shijiadong, Fengshun County; 9. Meiguang Village, Mt Lianhua, Wuhua County, Meizhou City (type locality of H. glaucus syn. nov.). The map is derived from Tianditu (www.tianditu.gov.cn). Phylogenetic analyses In total, 76 samples were used for phylogenetic analyses, encompassing 11 newly-sequenced individuals (three of the Hypselotriton population from Mt Fenghuang, four of H. orphicus from Jiexi County and two from Fengshun Coun- ty in Guangdong and two of H. yunnanensis from Honghe County in Yunnan) and others obtained from GenBank. Genomic DNA was extracted, using a DNA extraction kit from Tiangen Biotech (Beijing) Co., Ltd. Two mitochondri- on genes, namely NADH dehydrogenase subunit 2 (VD2) and Cytochrome b (cytb) were amplified for phylogenetic analyses. Primers for ND2 were L4437 (5’-AAGCTTTC- GGGCCCATACC-3’) and 5081R (5’-GICGTAGGGT- CAAAGCCTGC-3’) and, for cytb, these were 14052F (5’-CCTGGGCTCTAACCAAGACC-3’) and 15293R (5’-TCGGCTTACAAGACCGATGT-3’). PCR amplifica- tions were processed with the cycling conditions of: initial denaturing step at 95 °C for 4 min, 35 cycles of denaturing at 95 °C for 40 s, annealing at 53 °C for 34 s and exten- sion at 72 °C for 60 s and a final extension step at 72 °C for 10 min. PCR products were purified with spin columns and then sequenced with both forward and reverse primers using BigDye Terminator Cycle Sequencing Kit from Ap- plied Biosystems, on an ABI Prism 3730 automated DNA sequencer by Guangzhou Jierui Biotechnology Co., Ltd. All sequences were deposited in GenBank (Table 1). For phylogenetic analyses, DNA sequences were aligned by the Clustal W algorithm with default pa- rameters (Thompson et al. 1997). PartitionFinder2 was used to test the best partitioning scheme and jModelTest v.2.1.2 was used to test the best fitting nucleotide substi- tution model. Bayesian Inference (BI) in MrBayes 3.2.4 (Ronquist et al. 2012) and Maximum Likelihood (ML) in RAxML GUI (Silvestro and Michalak 2012) were used to conduct phylogenetic analyses. For the ML anal- ysis, an optimal tree was obtained and branch supports were evaluated with 1000 rapid bootstrapping replicates. zse.pensoft.net 1124 Wang, J. et al.: A new species and a new subgenus of Hypselotriton Table 1. Localities, voucher information and GenBank accession numbers for all samples used in this study. ID Genus / Species Hypselotriton oolong sp. Hypselotriton oolong sp. Hypselotriton oolong sp. Hypselotriton oolong sp. Hypselotriton oolong sp. Hypselotriton oolong sp. Hypselotriton oolong sp. nov. nov. nov. nov. nov. nov. nov. Hypselotriton sp. Hypselotriton sp. 10 ~—_Hypselotriton sp. 11 ~~ Hypselotriton sp. 12 ~~ Hypselotriton orphicus 13. -Hypselotriton orphicus 14 ~~ Hypselotriton orphicus 15 ~~ Hypselotriton orphicus 16 ~—-Hypselotriton orphicus 17 ~~ Hypselotriton orphicus 18 — Hypselotriton glaucus syn. nov. 19 ~—_Hypselotriton glaucus syn. nov. 20 ~~ _Hypselotriton cyanurus 21 ~~ _Hypselotriton cyanurus 22 — _Hypselotriton cyanurus 23 ~~ -Hypselotriton fudingensis 24 — Hypselotriton fudingensis 25 — Hypselotriton fudingensis 26 ~—-Hypselotriton jiaoren comb. nov. 27 Hypselotriton jiaoren comb. nov. 28 ~~ Hypselotriton jiaoren comb. nov. WO OND OF WN FE 29 ~——-Hypselotriton maguae comb. nov. 30 ~——-Hypselotriton maguae comb. nov. Si Hypselotriton orientalis 32 ~~ Hypselotriton orientalis 33. ~~ Hypselotriton orientalis 34 ~~ Hypselotriton orientalis 35 —_Hypselotriton orientalis 36 ~——-Hypselotriton orientalis 37 — Hypselotriton orientalis 38 ~~ Hypselotriton orientalis 39 ~—— Hypselotriton orientalis 40 ~=——-Hypselotriton orientalis 41 Hypselotriton yunnanensis 42 — Hypselotriton yunnanensis 43 Hypselotriton yunnanensis 44 ~~ Hypselotriton yunnanensis 45 Hypselotriton yunnanensis 46 ~~ -Hypselotriton yunnanensis 47 — Hypselotriton yunnanensis 48 — Cynops pyrrhogaster 49 = Cynops pyrrhogaster 50 Cynops pyrrhogaster 51 _Laotriton laoensis 52 — Pachytriton airobranchiatus 53. Pachytriton archospotus 54 — Pachytriton brevipes 55 ~Pachytriton feli 56 ~—_-Pachytriton granulosus 57 Pachytriton inexpectatus 58 — Pachytriton moi 59 — Pachytriton wuguantui 60 ~=Pachytriton xanthospilos 61 Paramesotriton aurantius 62 Paramesotriton caudopunctatus 63 Paramesotriton chinensis zse.pensoft.net Voucher CIB 121429 CIB 121430 SYS a009274 KIZ 09816 KIZ 09819 KIZ 09820 KIZ 09821 KIZ 09839 KIZ 09843 KIZ 09905 KIZ 09908 GEP a008 GEP a009 GEP a010 GEP a011 GEP a314 GEP a315 KIZ 09793 KIZ 09799 KIZ 02330 KIZ 02331 KIZ 02332 KIZ 012918 KIZ 012214 KIZ 012217 SYS a008787 SYS a008788 SYS a008789 CIB 118535 SYS a007032 KIZ 06358 CIB 97919 KIZ 012940 MVZ 230345 CIB 97867 KIZ 013017 YPX25002 KIZ 020536 KIZ 021962 KIZ 021844 CIB 121432 CIB 121433 SYS a007780 KIZ 01445 KIZ 021922 KIZ 022157 KIZ 022161 KIZ 09755 KIZ 09757 MVZ 263718 FMNH 255452 SWUF YZY0301 CIB 95950 KIZ 08926 KIZ 04228 KIZ 012977 KIZ 05203 KIZ 07767 KIZ 08756 KIZ 06750 KIZ 012879 KIZ 03903 KIZ 013010 China: China: China: China: China: China: China: Guangdong Guangdong Guangdong Guangdong Guangdong Guangdong Guangdong Locality : Chaozhou: : Chaozhou: : Chaozhou: : Chaozhou: : Chaozhou: : Chaozhou: : Chaozhou: Mt Fenghuang Mt Fenghuang Mt Fenghuang Mt Fenghuang Mt Fenghuang Mt Fenghuang Mt Fenghuang China: Fujian: Dehua: Mt Daiyun China: Fujian: Dehua: Mt Daiyun China: Fujian: Yongtai China: Fujian: Yongtai China: Guangdong: Jiexi: Mt Dabei China: Guangdong: Jiexi: Mt Dabei China: Guangdong: Jiexi: Mt Dabei China: Guangdong: Jiexi: Mt Dabei China: Guangdong: Fengshun: Mt Shijiadong China: Guangdong: Fengshun: Mt Shijiadong China: Guangdong: Wuhua: Mianyang China: Guangdong: Wuhua: Mianyang China: Guizhou: Shuicheng China: Guizhou: Shuicheng China: Guizhou: Shuicheng China: Fujian: Zherong China: Fujian: Ningde: Qingyu China: Fujian: Ningde: Qingyu China: Guangdong: Yingde China: Guangdong: Yingde China: Guangdong: Yingde China: Jiangxi: Nancheng: Mt Magu China: Jiangxi: Nancheng: Mt Magu China: Zhejiang: Jinhua China: Zhejiang: Quzhou China: Zhejiang: Tiantai China: Zhejiang: Hanzhou China: Jiangxi: Wannian China: Henan: Xinyang: Mt Jigong China: Jiangxi: Wuyuan China: Jiangxi: Jiujiang China: Anhui: Xiuning China: Anhui: Huoshan China: Yunnan: Honghe China: Yunnan: Honghe China: Yunnan: Shiping China: Yunnan: Ning’er China: Yunnan: Chuxiong: Zijing China: Yunnan: Kunming: Huahongdong China: Yunnan: Kunming: Gulu Japan Japan Japan Laos China: Guangdong: Huizhou: Mt Lianhua China: Jiangxi: Ganzhou: Mt Qiyun China: Fujian: Nanping: Mt Wuyi China: Anhui: Huangshan: Mt Huangshan China: Zhejiang: Taizhou: Tiantai China: Guangxi: Laibin: Mt Dayao China: Guangxi: Guilin: Mt Mao’er China: Guangxi: Hezhou China: Hunan: Chenzhou: Mt Mangshan China: Fujian: Ningde: Zherong China: Guizhou: Qiandongnan: Mt Leigong China: Zhejiang: Ningbo: Baixi ND2 PP987004 PP987005 PP987003 ON793719 ON793720 ON793721 ON793722 ON793723 ON793725 ON793728 ON793730 PP986999 PP987000 PP987001 PP987002 PP987008 PP987009 ON793715 ON793716 / ON793754 ON793755 ON793745 ON793743 ON793744 0Q116680 0Q116681 0Q116682 0Q116685 0Q116686 ON793718 GU301790 ON793731 EU880311 GU301788 ON793734 ON793740 ON793738 ON793737 ON793742 PP987006 PP987007 0Q116687 ON793756 ON793749 ON793751 ON793753 ji / EU880313 EU880328 MG732934 GQ303629 ON793903 ON793879 ON793920 ON793885 ON793895 ON793901 ON793907 ON794099 ON794069 ON794108 Cytb PP987015 PP987016 PP987014 ON793672 ON793673 ON793674 ON793675 ON793676 ON793678 ON793681 ON793683 PP987010 PP987011 PP987012 PP987013 PP987019 PP987020 ON793668 ON793669 ON793711 ON793712 ON793713 ON793698 ON793696 ON793697 / / / ON793671 / ON793684 EU880311 / ON793687 ON793694 ON793692 ON793691 ON793695 PP987017 PP987018 / ON793714 ON793706 ON793708 ON793710 ON793699 ON793701 EU880313 EU880328 MG732932 GQ303666 ON793838 ON793813 ON793856 ON793819 ON793830 ON793836 ON793842 ON794033 ON794003 ON794042 Zoosyst. Evol. 100 (3) 2024, 1121-1134 ID Genus / Species Voucher 64 ~~ Paramesotriton deloustali MVZ 223629 65 = Paramesotriton fuzhongensis KIZ 08568 66 Paramesotriton guangxiensis KIZ 09285 67 ~=Paramesotriton hongkongensis KIZ 01577 68 Paramesotriton labiatus KIZ 08769 69 Paramesotriton longliensis KIZ 03343 70. ~=Paramesotriton maolanensis GZNU 2006030004 71 ~~ Paramesotriton gixilingensis KIZ 022289 72. Paramesotriton wulingensis KIZ 03102 73. Paramesotriton yunwuensis KIZ 09676 74 Paramesotriton zhijinensis KIZ YPX6178 75 Calotriton asper Vieites 01 76 ~~ Euproctus platycephalus DBW-MVZ 01 N25 Locality ND2 Cytb Vietnam: Tam Dao: Yinh Yen FJ744600 GQ303669 China: Guangxi: Hezhou: Chaodong ON794073 ON794007 China: Guangxi: Chongzuo: Ningming ON794071 ON794005 China: Hong Kong: Tai Mo Shan ON794111 ON794045 China: Guangxi: Laibin: Mt Dayao ON794077 ON794011 China: Guizhou: Qiannan: Longli ON794064 ON793998 China: Guizhou: Qiannan: Libo JF438993 JX480887 China: Jiangxi: Jian: Mt Qixiling ON794120 ON794055 China: Chongqing: Youyang ON794123 ON794058 China: Guangdong: Yunfu: Luoding ON794086 ON794020 China: Guizhou: Bijie: Zhijing ON794067 ON794001 NA EU880307 EU880307 NA EU880317 EU880317 For the BI analysis, two independent runs with four Mar- kov Chain Monte Carlo simulations were performed for ten million iterations and sampled every 1000 iterations. The first 25% of the samples were discarded as burn-in after the standard deviation of split frequencies of the two runs was less than a value of 0.01. The remaining trees were used to create a consensus tree. Results The BI and ML analyses resulted in identical topologies (Fig. 2). Most major nodes were well supported with the Bayesian posterior probabilities (BPP) > 0.95 and the ML bootstrap supports (BS) > 70. All samples of Hypselo- triton formed a monophyletic group, as the sister taxon to the monophyletic group including Paramesotriton, Pachytriton and Laotriton, but paraphyletic with Cynops. This is corresponding to previous studies (Rancilhac et al. 2021; Yuan et al. 2022), supporting the resurrection of Hypselotriton as a distinct genus. Within Hypselotriton, three distinct and divergent clades were revealed, as those recovered in Lyu et al. (2023a). Morphological comparisons further sustained the differences amongst these clades (present below), which are treated as representing three subgenera. The clade involving H. orientalis has been proposed as sub- genus Cynotriton and the clade involving H. cyanurus is considered to represent the nominotypical subgenus Hypselotriton (Dubois and Raffaélli 2011; Raffaélli 2022). A new subgenus is nominated tn this work to ac- commodate species within the remaining unnamed clade. As shown in the tree, the Hypselotriton populations dis- tributed in eastern Guangdong have been separated into two distinct and distant lineages. Topotypical samples of H. orphicus from Jiexi were clustered with samples of type series of H. glaucus in a lineage and almost without genetic divergences, indicating these samples should be conspe- Key to subgenus within Hypselotriton cific. Morphological examination further confirmed such insight (present below). This lineage was the sister taxon to H. Jiaoren comb. nov. from northern Guangdong and col- lectively constituted the new subgenus as mentioned above. Samples of Hypselotriton population from Mt Feng- huang, which was employed as H. orphicus (Yuan et al. 2013, 2022; Lyu et al. 2023a), formed an independent lineage within the subgenus Cynotriton, representing the southernmost lineage of this subgenus (Fig. 1). Speci- mens of this lineage showed distinct differences in mor- phology that can be distinguished from H. orphicus, as well as all known congeners of Hypselotriton. Thus, this lineage represents an unnamed species that is described in this work. Moreover, the samples of Hypselotriton pop- ulation from central Fujian, which was also employed as H. orphicus (Yuan et al. 2013, 2022; Lyu et al. 2023a), formed a distinct lineage as the sister taxon to the Mt Fen- ghuang lineage, with small genetic divergence. The tax- onomic status of this lineage is further remarked below. Systematics Genus Hypselotriton Wolterstorff, 1934 Type species. Molge wolterstorffi Boulenger, 1905, by original designation. Common name. Chinese Fire-bellied Newts (in En- glish) / dian yuan sht (WAY in Chinese). Distribution. Endemic in mainland China. Remark. In spite of the paraphyletic relationships and distinct geographical isolation, species of the mainland genus Hypselotriton are very similar to those of insular genus Cynops in morphology, resulting in the prolonged categorising of these Fire-bellied Newts in a single genus. Hypselotriton can be distinguished from Cynops by the inconspicuous parotoid gland (vs. well developed) and smooth or slightly granular skin (vs. distinctly granular). 1 Distinct orange-red spot behind the eye present; blue lateral stripe on tail in breeding males present........ Hypselotriton - Distinct orange-red spot behind the eye absent; blue lateral stripe on tail in breeding males absent ..............cccceeee eee 2 zse.pensoft.net 1126 Wang, J. et al.: Anew species and a new subgenus of Hypselotriton 2 Mid-dorsal vertebral ridge continuous; irregular greyish-blue patches on dorsum and lateral tail in breeding males and PEL SS ORO SOM Lt tetpe Mee Maat: 25.6 6 brie Me aS ee nda ei nated att nceenit cate ata ile nbeknphepsedenaiangals te Hakkatriton subgen. nov. - Mid-dorsal vertebral ridge interrupted; irregular greyish-blue patches on dorsum and lateral tail in breeding males and Io) w ast Koka | olet =i gt AAR SR: SCROLL Re ON 202 gee Pe SD) ee ETON OS OS ote Cynotriton Subgenus Hypselotriton Wolterstorff, 1934 Type species. Molge wolterstorffi Boulenger, 1905, by original designation. Diagnosis. (1) small to large size; (2) gular fold pres- ent; (3) mid-dorsal vertebral ridge continuous; (4) meta- carpal and metatarsal tubercles on external side of hands and feet present; (5) postocular orange spot present; (6) blue lateral stripe on tail in breeding males present. Content and remarks. This subgenus includes all known Fire-bellied Newts populations distributed in the Yunnan-Guizhou Plateau in southwestern China. Currently, four nomenclatures have been provisionally documented, i.e. H. (Hy.) chenggongensis (Kou & Xing, 1983), H. (Hy.) cyanurus (Liu, Hu & Yang, 1962), H. (Hy.) wolterstorffi (Boulenger, 1905) and H. (Hy.) yunnanensis (Yang, 1979) (Lyu et al. 2023a). However, the exact taxonomic status for these populations/species is still unresolved (Raffaélli 2022; Lyu et al. 2023a). Thus, it 1s improper to present a key to these nomenclatures before a comprehensive study to precisely delimitate their taxonomic placements. Subgenus Hakkatriton subgen. nov. https://zoobank.org/504DCCDF-32D2-4894-AFBA-5766752AD418 Type species. Cynops orphicus Risch, 1983, by present designation. Etymology. The nomen of Hakkatriton subgen. nov. is derived from Hakka, referring to its distribution in northern and eastern Guangdong where is the settle- ment of Hakka people and generic nomen Triton Lau- renti, 1768. Diagnosis. (1) small size; (2) gular fold absent; (3) mid-dorsal vertebral ridge continuous; (4) metacarpal and metatarsal tubercles on external side of hands and feet absent; (5) postocular orange spot absent; (6) blue lateral stripe on tail in breeding males absent; (7) irregular grey- ish-blue patches on dorsum and lateral tail in breeding males and females present. Content. Two species distributed in isolated regions in northern and eastern Guangdong in southern China, respectively (Fig. 1). Key to species of the subgenus Hakkatriton subgen. nov. 1 Ventral forearms and shanks uniformly dark brown; tail Uniformly COIOULE ........... cee eecceeee eee eeeeeeee eee eeeees H. (Ha.) Jiaoren ~ Ventral forearms and shanks with bright orange patches; tail with irregular black spots ................:0000 H. (Ha.) orphicus Hypselotriton (Hakkatriton) orphicus (Risch, 1983) Pachytriton brevipes (Sauvage, 1876) — Pope and Boring (1940); Gressitt (1941) Cynops shataukokensis Freytag & Eberhardt, 1977 — Freytag (1979) Cynops orphicus Risch, 1983 — Risch (1983); Fei et al. (2006, 2012); Fei and Ye (2016) Hypselotriton (Pingia) orphicus — Dubois and Raffaélli (2009) Hypselotriton (Cynotriton) orphicus — Dubois and Raffaélli (2011) Cynops glaucus Yuan, Jiang, Ding, Zhang & Che, 2013 syn. nov. — Yuan et al. (2013, 2022); Fei and Ye (2016); Lyu et al. (2023a) Hypselotriton (Cynotriton) glaucus — Raffaélli (2022) Type. Holotype. Cutna * 3; Guangdong Province, Shan- tou Region [now belonging to Jieyang City, Jiexi Coun- ty], Tai-Yong [Dayang Township]; 23°35'N, 115°S1'E [=23.58°N, 115.85°E; located in Liangtian Township, see remarks below], 640 m elev.; 4 Aug. 1936; JL Gressitt leg.; MVZ 22474. Paratypes. CHINA * 97 adult specimens; same data as for holotype; MVZ 22416—73, 22475—506 [90 speci- mens], MNHN 1980.4096—4098 [3 specimens, formerly MVZ 24134—36], AMNH 46174, CAS 78704, 2 unnum- zse.pensoft.net bered specimens in the Department of Biology, Yenching University [probably lost according to Risch (1983)]. Examined specimens. CHINA * 463; Guang- dong Province, Jieyang City, Jiexi County, Mt Dabei; 23.55°N, 115.89°E, ca. 490 m elev.; GEP a008, 010— 011, CIB 121434 * 299; same data as for preceding; GEP a009, CIB 121435 * 24'4; Guangdong Province, Meizhou City, Fengshun County, Mt Hongtuzhang; 23.78°N, 115.96°E, ca. 1200 m elev.; GEP a263—264 « 343; Guangdong Province, Meizhou City, Fengshun County, Mt Shijiadong; 23.81°N, 116.33°E, ca. 1140 m elev.; GEP a314—316 * 39 9; same data as for preceding; GEP a317-319. Referred specimens. Seven specimens labelled as “Cynops glaucus” in Lyu et al. (2023a). CHINA * 299; Guangdong Province, Jieyang City, Jiexi County, Mt Dabei; SYS a000729, 8511 * 14; Guangdong Province, Jieyang City, Jiexi County, Mt. Liwangzhang; 23.64°N, 115.81°E, ca. 990 m elev.; SYS a008602 * 19; same data as for preceding; SYS a008601 * 29 9: Guangdong Prov- ince, Meizhou City, Fengshun County, Mt. Tongguzhang; 24.18°N, 116.35°E, ca. 1500 m elev.; SYS a000730, 4743 ¢ 19; same data as for preceding; SYS a000731. Zoosyst. Evol. 100 (3) 2024, 1121-1134 +/+ Cynotriton +/+ +/+ Hypselotriton an +/+ Hypselotriton +/+ +/+ Hakkatriton subgen. nov. +/+ +/+ -/- +/+ a +/+ +/+ +/+ +/F +/+ +/+ 0.02 —— +/+ +/+ +/+ +t +/+] 70 +/+ +/+} | -/ +/+ +} a +/+p pyrrhogaster +/+ pyrrhogaster pyrrhogaster +/+ +/+ 1127 (1) Mt Fenghuang, Guangdong (2) Mt Fenghuang, Guangdong (3) Mt Fenghuang, Guangdong (4) Mt Fenghuang, Guangdong (5) Mt Fenghuang, Guangdong 4/4 (6) Mt Fenghuang, Guangdong (7) Mt Fenghuang, Guangdong (8) Mt Daiyun, Fujian (9) Mt Daiyun, Fujian +/+], (10) Yongtai, Fujian (11) Yongtai, Fujian +/+, (29) Mt Magu, Jiangxi (30) Mt Magu, Jiangxi +/+ oolong sp. nov. sp. maguae comb. nov. (23) Zherong, Fujian (24) Ningde, Fujian (25) Ningde, Fujian (31) Jinhua, Zhejiang (33) Tiantai, Zhejiang (32) Quzhou, Zhejiang (35) Wannian, Jiangxi (38) Jiujiang, Jiangxi (39) Xiuning, Anhui (40) Huoshan, Anhui (37) Wuyuan, Jiangxi (34) Hanzhou, Zhejiang (36) Mt Jigong, Henan (41) Honghe, Yunnan (42) Honghe, Yunnan (43) Shiping, Yunnan (45) Chuxiong, Yunnan (46) Kunming, Yunnan (47) Kunming, Yunnan (44) Ning'er, Yunnan (20) Shuicheng, Guizhou (21) Shuicheng, Guizhou (22) Shuicheng, Guizhou (26) Yingde, Guangdong (27) Yingde, Guangdong (28) Yingde, Guangdong (19) Wuhua, Guangdong (18) Wuhua, Guangdong (13) Jiexi, Guangdong (14) Jiexi, Guangdong +/+ (12) Jiexi, Guangdong (15) Jiexi, Guangdong (16) Fengshun, Guangdong (17) Fengshun, Guangdong aurantius hongkongensis chinensis gixilingensis deloustali fuzhongensis guangxiensis yunwuensis labiatus caudopunctatus wulingensis longliensis maolanensis zhijinensis fudingensis orientalis yunnanensis cyanurus +/+ jiaoren comb. nov. glaucus syn. nov. (type series) orphicus (topotypes) Paramesotriton airobranchiatus wuguanfui xanthospilos brevipes et Pachytriton granulosus archospotus inexpectatus moi laoensis ] Laotriton Cynops Euproctus platycephalus Calotriton asper Figure 2. Bayesian Inference and Maximum-Likelihood phylogenies, based on mitochondrial ND2 and cytb genes. Bayesian poste- rior probabilities (BPP) > 0.95 and the bootstrap supports (BS) > 70 are labelled with “+” and otherwise labelled with “-”. Etymology. The specific name orphicus is derived from the Greek legendary musician and poet Orpheus, in memo- ry of several people who passed away in 1982 (Risch 1983). Common name. Dayang Fire-bellied Newt (in En- glish) / chao shan rong yuan (37/4245 in Chinese). Revised diagnosis. (1) small body size, TOL 68.5—77.0 mm in adult males, 85.1—-99.7 mm in adult females; (2) parotoid gland inconspicuous; (3) postocular orange spot absent; (4) surface smooth, finely granulated, gular fold absent; (5) continuous vertebral ridge weak and inconspic- uous; (6) fingers and toes overlapping when fore-limb and hind-limb adpressed towards each other along body; (7) ground colour dark brown to olive brown, with irregular greyish-blue patches on dorsum and lateral tail in breed- ing males and females; (8) lateral tail with black spots; (9) ground colour of venter dark brown with irregular bright or- ange patches, bright orange blotches on chin, ventral limbs and cloaca; (10) ventral tail with a bright orange stripe. Description of new specimens. Body slender and small-sized, TOL 68.5—77.0 mm in adult males, 85.1— 99.7 mm in adult females, with detailed measurements listed in Table 2. Head oval in dorsal view; snout trun- cate, projecting slightly beyond mandible; nostril small, but conspicuous; labial fold developed on posterior part of upper jaw; tongue elongate, enlarged anteriorly, with free lateral margin; vomerine tooth patch “””’-shaped:; eye small, not extending beyond lateral margins of head; an inconspicuous longitudinal ridge found posterior to each eye; parotoid gland inconspicuous, gill remnants absent; gular fold absent. zse.pensoft.net 1128 Wang, J. et al.: A new species and a new subgenus of Hypselotriton Table 2. Measurements (in mm) of Hypselotriton (Hakkatriton) orphicus and H. (Cynotriton) oolong sp. nov. Voucher Sex TOL SVL TAL TAD HL HW Hypselotriton (Hakkatriton) orphicus GEP a008 M 73:5 41.9 29.1 6.1 Syl 8.8 GEP a010 M F335 43.3 82.3 5.0 12) 8.8 GEP a011 M TAO 44.2 328 Or 12.8 9.7 GEP a263 M 75.6 44,3 SURES: 4.8 10.0 8.6 GEP a264 M 15.9 43.0 32.9 4.9 12.6 8.7 GEP a314 M 72.1 42.3 29.8 6.6 12.5 10.0 GEP a315 M 70.7 40.8 29.9 6.4 11.8 9.4 GEP a316 M 74.5 42.2 3283 6.2 12.2. 9.3 CIB 121434 M 68.5 41.3 2782 6.7 11.9 8.7 SYS a000730 M 70.7 43.4 nae aes 6.8 Lee. 9.2 SYS a004743. M 79.8 49.9 29.9 6.8 L22. 10.4 SYS a008602 -M 135 43.5 30.0 6.4 12.5 10.0 GEP a009 F 85.1 48.7 36.4 6.6 14.1 9.9 GEP a317 F 86.1 48.1 38.1 6.1 13.6 10.8 GEP a318 F 91.2 50.3 40.9 7.4 14.6 10.8 GEP a319 F 91.0 50.6 40.4 6.3 14.2 10.7 CIB 121435 F 88.9 48.6 40.3 St 13:5 10:3 SYS a000729 OF 87.5 50.2 #3 5.9 1337, 10.1 SYS a000731 SOF 88.6 49.6 39.0 6.9 14.1 11.4 SYS a008511 —SO#F 90.2 50.6 39.6 57 13:7 10.5 SYS a008601 =F 99:7 S516 44.1 “5 14.8 11.6 Hypselotriton (Cynotriton) oolong sp. nov. CIB 121430 M 70.7 45.2 25,5 6.1 12.5 9.1 CIB 121429 M 69.9 43.0 26.9 Sz 12.4 8.6 SYS a009274. M 70.0 43.5 26.5 5.0 11.6 8.4 Surface smooth, finely granular; a few inconspicuous longitudinal wrinkles present on chin; continuous verte- bral ridge weak and inconspicuous; cloacal opening oval, slightly protruding. Limbs slender, fingers and toes overlapping when fore- limb and hind-limb adpressed towards each other along body; four fingers and five toes, slender and elongated, lack webbing; relative length of fingers I < IV < II < III; relative length of toes |< V < I]