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Tylototriton tongziensis sp. nov. Holotype. Figs. 3, 4. Adult male, CIB WB2020190, collected on 21 May 2020 by Shize Li from Huanglian Nature Reserve (28° 29′ N, 107° 02′ E, ca. 1510 m a.s.l.), Tongzi County, Guizhou Province, China. Paratypes. Three adult males, CIB WB2020081511 collected on 15 August 2020 by Jing Liu, CIB WB2020191 on 21 May 2020 by Bin Wang, and CIB TZ20160714005 collected on 14 July 2016 by Shize Li; two adult females, CIB TZ20160714010 and CIB TZ20160714002 collected on July 2016 by Shize Li; two larvae, CIB WB2020201 and CIB WB2020202 on 21 May 2020 by Shengchao Shi. All paratypes were collected from the same location as the holotype. Diagnosis: Tylototriton tongziensis sp. nov. could be distinguished from its congeners by a combination of the following characters: (1) body size medium (TOL 120.5–135.1 mm, SVL 61.1–65.9 mm in males, and TOL 123.5–127.6 mm, SVL 66.7–69.2 mm in females); (2) gular fold present; (3) the tail length shorter than the snoutvent length; (4) the distal ends and ventral surfaces of digits, peripheral area of cloaca, and the lower margin of tail orange; (5) the distal tips of the limbs greatly overlapping when the fore and hind limbs being pressed along the trunk; (6) fingertips reaching to the level beyond the snout when the forelimbs being stretched forward; (7) nodulelike warts on body sides continuous and no obvious. Description of holotype. SVL 62.9 mm; TOL 131.4 mm; head slightly longer than wide (HDL/HDW = 1.02); head slightly concave on the top; snout nearly square, extending beyond the lower lip; naris nearly snout; snout truncate in dorsal view; fabial fold absent; dorsolateral bony ridges on head prominent, moderately protruding, from above snout to above anterior end of parotoid, posterior ends relatively thick and scrolled inside; mid-dorsal ridge on head distinct and thin; parotoids enlarged, projecting backwards; tinter-nasal space is smaller than the inter-orbital space; eyes protrude from the dorsolateral portion of the head; the oral fissure flat; the joint of jaw articulation lies posterior to the caudal margin of the eyes; fine teeth present on the edge of jaw, vomerine teeth long and prominent, form a “∧” shape; tongue oval and nearly entirely fixed at bottom but free at both lateral edges; neck is rounded and thick, gular fold present. Body stout; skin on lateral body with no obvious presence of small glands, glandular vertebral ridge high, slightly rough and segmented, extending from top of head to base of tail, rib nodules indistinct, with similar sizes. Four limbs relatively slender, fingers from fore-and hind limbs overlap when addressed along body; tips of fingers beyond snout when addressed along head; relative fingers length III> II> IV> I, and relative toes length III> IV> II> I> V; fingers and digits are with none webbed. The tail length shorter than the snout-vent length; the tail laterally compressed, thin and tip acuminated, distinctly thin and high; the cloaca long and narrow, and the cloacal region slightly bulbous. The skin rough, and body covered with small tubercles; the labial margin, distal limbs, ventral limbs and the ventral edge of the tail smooth; on the lateral dorsum of the body, tubercles small, appearing to form lines that extend from the shoulder to the base of the tail and the nodule-like tubercles continuously distributed with no clear separation between neighboring tubercles; the ventral tubercles dense and distinct. ......continued on the next page TABLE 4. (Continued) *: the population from Leishan County, Guizhou Province, China. Color in life. In life, the individual black or blackish-brown; the distal ends and ventral surface of digits, the peripheral area of cloaca, and the ventral edge of tail orange; the orange region between the ventral edge of the tail and the peripheral area of the cloaca connected (Fig. 3). Color in preservative. The specimen in preservative is blackish-brown. The orange coloration of the distal ends and ventral surface of digits, the peripheral area of the cloaca, and the ventral edge of the tail fades to lacte (Fig. 4). Secondary sexual characteristics. The female cloacal hole short and its inner cloacal wall have no papilla. The male has papilla on its inner cloacal wall and its cloaca is a long slit (Figs. 3H, 4F). Larva. Two larvae, body slender. The forelimbs and hindlimbs clearly fully grown up. Head longer than wide, eyes relatively large on the flank and three pairs of external gills having obtuse anterior ends, while the body and tail being laterally compressed. Skin smooth, fingers and toes transparent, body black-brown, tail black with small yellow spots. Costal grooves for muscles of larvae indistinctly. The dorsal fin fold comparatively tall and rises from the region between the posterior head and anterior body. The short and thick ventral fin fold of the tail extends from the cloaca to the end of the tail. Morphological comparisons. Tylototriton tongziensis sp. nov. is phylogenetically clustered into a clade V containing T. dabienicus, T. anhuiensis, T. broadoridgus, T. maolanensis, T. “ daloushanensis ”, T. wenxianensis, T. liuyangensis, and T. lizhengchangi, and could be identified from these closely related species on series of morphological characters (Table 5). Tylototriton tongziensis sp. nov. differs from T. dabienicus by having a shorter total length in female, TOL 123.5–127.6 mm (vs. 134.9–155.5 mm), finger tips beyond the snout when forelimbs stretched forward (vs. reaching anterior corner of eyes), and tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched). Tylototriton tongziensis sp. nov. differs from T. anhuiensis by tips of fore-and hindlimbs overlapping when addressed along body (vs. just meeting). Tylototriton tongziensis sp. nov. differs from T. broadoridgus by having a shorter total length in female, TOL 123.5–127.6 mm (vs. 138.9–162.5 mm); tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched); gular fold present (vs. absent). Tylototriton tongziensis sp. nov. differs from T. maolanensis by having a shorter total length, TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 151.0–172.0 mm in males and 142.7–170.5 mm in females); nodule-like warts on body sides continuous and no obvious (vs. separated and distinct). Tylototriton tongziensis sp. nov. differs from T. “ daloushanensis ” by having a shorter snout-vent length in females, SVL 66.7–69.2 mm (vs. 70.5–100.3 mm); finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level between eyes to nostrils). Tylototriton tongziensis sp. nov. differs from T. wenxianensis by gular fold present (vs. absent); tips of fore-and hindlimbs overlapping when addressed along body (vs. meeting or overlapping). Tylototriton tongziensis sp. nov. differs from T. liuyangensis by having a shorter total length in females, TOL 123.5–127.6 mm (vs. 138.6–154.2 mm); finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye); tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched). Tylototriton tongziensis sp. nov. differs from T. lizhengchangi by having a shorter total length, TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 145.6–173.0 mm in males and 150.0– 156.5 mm in females). For the species within clade III, Tylototriton tongziensis sp. nov. differs from T. panhai by the absence of orange markings on the parotoid (vs. present); from T. vietnamensis by the presence of gular fold (vs. absent). Tylototriton tongziensis sp. nov. further differs from T. pasmansi by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. sparreboomi by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of nostril), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. asperrimus by nodule-like warts on body sides continuous and no obvious (vs. separated), and tips of fore-and hindlimbs overlapping when addressed along body (vs. meeting or overlapping); from T. thaiorum by nodule-like warts on body sides continuous and no obvious (vs. separated); from T. notialis by nodule-like warts on body sides continuous and no obvious (vs. separated), and orange markings on the parotoid and the rib nodules absent (vs. present); from T. sini by having a shorter total length in females, TOL 123.5–127.6 mm (vs. 144.5 mm), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. hainanensis by having a shorter total length in males, TOL 120.5–135.1 mm (vs. 137.0–148.0 mm) by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye); from T. ziegleri by nodule-like warts on body sides continuous and no obvious (vs. separated), and vertebral ridge not segmented (vs. segmented). Tylototriton tongziensis sp. nov. differs from T. anguliceps, T. himalayanus, T. kachinorum, T. kweichowensis, T. ngarsuensis, T. panwaensis, T. phukhaensis, T. podichthys, T. pulcherrimus, T. shanjing, T. shanorum, T. umphangensis, T. uyenoi, T. verrucosus, and T. yangi by having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail (vs. having distinct yellow to reddish brown markings on head, trunk, and tail edges in the latter). Tylototriton tongziensis sp. nov. differs from T. taliangensis by having a shorter total length TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 186.0–220.0 mm in males and 194.0–230.0 mm in females) and orange markings on the parotoid absent (vs. present); from T. pseudoverrucosus by having a shorter total length TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 156.2–173.0 mm in males and 178.2 mm in female), nodule-like warts on body sides continuous and no obvious (vs. separated), and having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail (vs. distinct yellow to reddish brown markings on head, trunk, and tail edges). Distribution. Tylototriton tongziensis sp. nov. is currently known from Huanglian Nature Reserve, Tongzi County, Guizhou Province, China. ......continued on the next page TABLE 5. (Continued) Ecology. The newts inhabit a mountain stream (Fig. 6) covered by evergreen broadleaf forest, at elevations of about 1530 m. Adult individuals can be observed in a small pond with muddy bottoms from early May to mid-August. The larvae were also found in the same pond from May to August, the water of pond was clear, and 3 m long and ca 2 m wide with water depth ca. 20 cm. Other syntopic species of amphibians recorded at the type locality include: Zhangixalus omeimontis (Stejneger, 1924), Z. chenfui (Liu, 1945), Hyla annectans (Jerdon, 1870), and Rana omeimontis Ye & Fei, 1993. Conservation recommendation. Tylototriton tongziensis sp. nov. is to date known only from a single location in Huanglian Nature Reserve, Tongzi County, Guizhou Province, China. The type locality is suffering from the disturbs of the development of tourism, and the population of the species is small according to our surveys. We recommend Tylototriton tongziensis sp. nov. to be listed as Critically Endangered (CR) [IUCN Red List criteria B1ab(iii)+2ab(iii)]. Etymology. The specific name tongziensis is a Latinize toponymic adjective that refers to Tongzi County, Guizhou Province, China, where the new species was firstly found. For the common name, we suggest Tongzi Knobby Newt (English) and Tong Zi You Yuan (Chinese)., Published as part of Li, Shi-Ze, Liu, Jing, Shi, Sheng-Chao, Wei, Gang & Wang, Bin, 2022, Description of a new species of the newt genus Tylototriton sensu lato (Amphibia: Urodela: Salamandridae) from southwestern China, pp. 248-268 in Zootaxa 5128 (2) on pages 254-264, DOI: 10.11646/zootaxa.5128.2.5, http://zenodo.org/record/6479851, {"references":["Wang, B., Nishikawa, K., Matsui, M., Nguyen, T. Q., Xie, F., Li, C., Khatiwada, J. R., Zhang, B. W., Gong, D. J., Mo, Y. M., Wei, G., Chen, X. H., Shen, Y. H., Yang, D. D., Xiong, R. C. & Jiang, J. P. (2018) Phylogenetic surveys on the newt genus Tylototriton sensu lato (Salamandridae, Caudata) reveal cryptic diversity and novel diversification promoted by historical climatic shifts. PeerJ, 6, e 4384. https: // doi. org / 10.7717 / peerj. 4384"]}

Tylototriton tongziensis sp. nov. Holotype. Figs. 3, 4. Adult male, CIB WB2020190, collected on 21 May 2020 by Shize Li from Huanglian Nature Reserve (28° 29′ N, 107° 02′ E, ca. 1510 m a.s.l.), Tongzi County, Guizhou Province, China. Paratypes. Three adult males, CIB WB2020081511 collected on 15 August 2020 by Jing Liu, CIB WB2020191 on 21 May 2020 by Bin Wang, and CIB TZ20160714005 collected on 14 July 2016 by Shize Li; two adult females, CIB TZ20160714010 and CIB TZ20160714002 collected on July 2016 by Shize Li; two larvae, CIB WB2020201 and CIB WB2020202 on 21 May 2020 by Shengchao Shi. All paratypes were collected from the same location as the holotype. Diagnosis: Tylototriton tongziensis sp. nov. could be distinguished from its congeners by a combination of the following characters: (1) body size medium (TOL 120.5–135.1 mm, SVL 61.1–65.9 mm in males, and TOL 123.5–127.6 mm, SVL 66.7–69.2 mm in females); (2) gular fold present; (3) the tail length shorter than the snoutvent length; (4) the distal ends and ventral surfaces of digits, peripheral area of cloaca, and the lower margin of tail orange; (5) the distal tips of the limbs greatly overlapping when the fore and hind limbs being pressed along the trunk; (6) fingertips reaching to the level beyond the snout when the forelimbs being stretched forward; (7) nodulelike warts on body sides continuous and no obvious. Description of holotype. SVL 62.9 mm; TOL 131.4 mm; head slightly longer than wide (HDL/HDW = 1.02); head slightly concave on the top; snout nearly square, extending beyond the lower lip; naris nearly snout; snout truncate in dorsal view; fabial fold absent; dorsolateral bony ridges on head prominent, moderately protruding, from above snout to above anterior end of parotoid, posterior ends relatively thick and scrolled inside; mid-dorsal ridge on head distinct and thin; parotoids enlarged, projecting backwards; tinter-nasal space is smaller than the inter-orbital space; eyes protrude from the dorsolateral portion of the head; the oral fissure flat; the joint of jaw articulation lies posterior to the caudal margin of the eyes; fine teeth present on the edge of jaw, vomerine teeth long and prominent, form a “∧” shape; tongue oval and nearly entirely fixed at bottom but free at both lateral edges; neck is rounded and thick, gular fold present. Body stout; skin on lateral body with no obvious presence of small glands, glandular vertebral ridge high, slightly rough and segmented, extending from top of head to base of tail, rib nodules indistinct, with similar sizes. Four limbs relatively slender, fingers from fore-and hind limbs overlap when addressed along body; tips of fingers beyond snout when addressed along head; relative fingers length III> II> IV> I, and relative toes length III> IV> II> I> V; fingers and digits are with none webbed. The tail length shorter than the snout-vent length; the tail laterally compressed, thin and tip acuminated, distinctly thin and high; the cloaca long and narrow, and the cloacal region slightly bulbous. The skin rough, and body covered with small tubercles; the labial margin, distal limbs, ventral limbs and the ventral edge of the tail smooth; on the lateral dorsum of the body, tubercles small, appearing to form lines that extend from the shoulder to the base of the tail and the nodule-like tubercles continuously distributed with no clear separation between neighboring tubercles; the ventral tubercles dense and distinct. ......continued on the next page TABLE 4. (Continued) *: the population from Leishan County, Guizhou Province, China. Color in life. In life, the individual black or blackish-brown; the distal ends and ventral surface of digits, the peripheral area of cloaca, and the ventral edge of tail orange; the orange region between the ventral edge of the tail and the peripheral area of the cloaca connected (Fig. 3). Color in preservative. The specimen in preservative is blackish-brown. The orange coloration of the distal ends and ventral surface of digits, the peripheral area of the cloaca, and the ventral edge of the tail fades to lacte (Fig. 4). Secondary sexual characteristics. The female cloacal hole short and its inner cloacal wall have no papilla. The male has papilla on its inner cloacal wall and its cloaca is a long slit (Figs. 3H, 4F). Larva. Two larvae, body slender. The forelimbs and hindlimbs clearly fully grown up. Head longer than wide, eyes relatively large on the flank and three pairs of external gills having obtuse anterior ends, while the body and tail being laterally compressed. Skin smooth, fingers and toes transparent, body black-brown, tail black with small yellow spots. Costal grooves for muscles of larvae indistinctly. The dorsal fin fold comparatively tall and rises from the region between the posterior head and anterior body. The short and thick ventral fin fold of the tail extends from the cloaca to the end of the tail. Morphological comparisons. Tylototriton tongziensis sp. nov. is phylogenetically clustered into a clade V containing T. dabienicus, T. anhuiensis, T. broadoridgus, T. maolanensis, T. “ daloushanensis ”, T. wenxianensis, T. liuyangensis, and T. lizhengchangi, and could be identified from these closely related species on series of morphological characters (Table 5). Tylototriton tongziensis sp. nov. differs from T. dabienicus by having a shorter total length in female, TOL 123.5–127.6 mm (vs. 134.9–155.5 mm), finger tips beyond the snout when forelimbs stretched forward (vs. reaching anterior corner of eyes), and tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched). Tylototriton tongziensis sp. nov. differs from T. anhuiensis by tips of fore-and hindlimbs overlapping when addressed along body (vs. just meeting). Tylototriton tongziensis sp. nov. differs from T. broadoridgus by having a shorter total length in female, TOL 123.5–127.6 mm (vs. 138.9–162.5 mm); tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched); gular fold present (vs. absent). Tylototriton tongziensis sp. nov. differs from T. maolanensis by having a shorter total length, TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 151.0–172.0 mm in males and 142.7–170.5 mm in females); nodule-like warts on body sides continuous and no obvious (vs. separated and distinct). Tylototriton tongziensis sp. nov. differs from T. “ daloushanensis ” by having a shorter snout-vent length in females, SVL 66.7–69.2 mm (vs. 70.5–100.3 mm); finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level between eyes to nostrils). Tylototriton tongziensis sp. nov. differs from T. wenxianensis by gular fold present (vs. absent); tips of fore-and hindlimbs overlapping when addressed along body (vs. meeting or overlapping). Tylototriton tongziensis sp. nov. differs from T. liuyangensis by having a shorter total length in females, TOL 123.5–127.6 mm (vs. 138.6–154.2 mm); finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye); tips of fore-and hindlimbs overlapping when addressed along body (vs. not touched). Tylototriton tongziensis sp. nov. differs from T. lizhengchangi by having a shorter total length, TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 145.6–173.0 mm in males and 150.0– 156.5 mm in females). For the species within clade III, Tylototriton tongziensis sp. nov. differs from T. panhai by the absence of orange markings on the parotoid (vs. present); from T. vietnamensis by the presence of gular fold (vs. absent). Tylototriton tongziensis sp. nov. further differs from T. pasmansi by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. sparreboomi by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of nostril), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. asperrimus by nodule-like warts on body sides continuous and no obvious (vs. separated), and tips of fore-and hindlimbs overlapping when addressed along body (vs. meeting or overlapping); from T. thaiorum by nodule-like warts on body sides continuous and no obvious (vs. separated); from T. notialis by nodule-like warts on body sides continuous and no obvious (vs. separated), and orange markings on the parotoid and the rib nodules absent (vs. present); from T. sini by having a shorter total length in females, TOL 123.5–127.6 mm (vs. 144.5 mm), and nodule-like warts on body sides continuous and no obvious (vs. separated); from T. hainanensis by having a shorter total length in males, TOL 120.5–135.1 mm (vs. 137.0–148.0 mm) by finger tips beyond the snout when forelimbs stretched forward (vs. reaching the level of eye); from T. ziegleri by nodule-like warts on body sides continuous and no obvious (vs. separated), and vertebral ridge not segmented (vs. segmented). Tylototriton tongziensis sp. nov. differs from T. anguliceps, T. himalayanus, T. kachinorum, T. kweichowensis, T. ngarsuensis, T. panwaensis, T. phukhaensis, T. podichthys, T. pulcherrimus, T. shanjing, T. shanorum, T. umphangensis, T. uyenoi, T. verrucosus, and T. yangi by having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail (vs. having distinct yellow to reddish brown markings on head, trunk, and tail edges in the latter). Tylototriton tongziensis sp. nov. differs from T. taliangensis by having a shorter total length TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 186.0–220.0 mm in males and 194.0–230.0 mm in females) and orange markings on the parotoid absent (vs. present); from T. pseudoverrucosus by having a shorter total length TOL 120.5–135.1 mm in males and 123.5–127.6 mm in females (vs. 156.2–173.0 mm in males and 178.2 mm in female), nodule-like warts on body sides continuous and no obvious (vs. separated), and having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail (vs. distinct yellow to reddish brown markings on head, trunk, and tail edges). Distribution. Tylototriton tongziensis sp. nov. is currently known from Huanglian Nature Reserve, Tongzi County, Guizhou Province, China. ......continued on the next page TABLE 5. (Continued) Ecology. The newts inhabit a mountain stream (Fig. 6) covered by evergreen broadleaf forest, at elevations of about 1530 m. Adult individuals can be observed in a small pond with muddy bottoms from early May to mid-August. The larvae were also found in the same pond from May to August, the water of pond was clear, and 3 m long and ca 2 m wide with water depth ca. 20 cm. Other syntopic species of amphibians recorded at the type locality include: Zhangixalus omeimontis (Stejneger, 1924), Z. chenfui (Liu, 1945), Hyla annectans (Jerdon, 1870), and Rana omeimontis Ye & Fei, 1993. Conservation recommendation. Tylototriton tongziensis sp. nov. is to date known only from a single location in Huanglian Nature Reserve, Tongzi County, Guizhou Province, China. The type locality is suffering from the disturbs of the development of tourism, and the population of the species is small according to our surveys. We recommend Tylototriton tongziensis sp. nov. to be listed as Critically Endangered (CR) [IUCN Red List criteria B1ab(iii)+2ab(iii)]. Etymology. The specific name tongziensis is a Latinize toponymic adjective that refers to Tongzi County, Guizhou Province, China, where the new species was firstly found. For the common name, we suggest Tongzi Knobby Newt (English) and Tong Zi You Yuan (Chinese).