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A new species of the genus Tylototriton is described from Ngoc Linh Mountain, Kon Tum Province, in the Central Highlands of Vietnam based on integrative taxonomy, namely by combining molecular and morphological evidence. Tylototriton ngoclinhensis sp. nov. differs from all other congeners based on morphological data, allopatric distribution, and molecular divergence. In terms of genetic divergence, Tylototriton ngoclinhensis sp. nov. distinctly differs from the sister species T. panhai (6.77%) and from T. ngarsuensis (12.36%) based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. Tylototriton ngoclinhensis sp. nov. is a moderate sized and robust salamander species with large cephalic edges, parotoids, and vertebral ridge orange in coloration. The new taxon differs from its congeners by a combination of the following morphological characteristics: size medium (SVL 60.8–66.5 mm, TL 57.6–61.8 mm in males, and SVL 72.5–75.6 mm, TL 62.9–67.9 mm in females); head longer than wide; parotoids very prominent and enlarged, projecting backwards; tail length shorter than snout-vent length; vertebral ridge large, high and glandular in appearance; 14 large and distinct dorsolateral glandular warts; gular fold present; tips of fore and hind limbs overlapping when adpressed along the body; tips of fingers reaching between eye and nostril when foreleg is laid forward; dorsal surface and lateral sides of the head, upper and lower lips, dorsolateral glandular warts, vertebral ridge, the peripheral area of the cloaca and the ventral edge of the tail orange in coloration; the presence of a distinct black line extending from the posterior end of the eye towards the shoulder. Tylototriton ngoclinhensis sp. nov. is restricted to evergreen montane forests near water bodies on Ngoc Linh Mountain. We suggest that the new species should be classified as Endangered (EN) in the IUCN Red List. This new important discovery represents the eighth Tylototriton taxon described from Vietnam, and at the same time constitutes the southernmost distributional record for the whole genus in Asia.

Herein, we provided a morphological description combined with a phylogenetic analysis of the northernmost Thai Tylototriton population, which was found in Doi Pha Hom Pok National Park (DPHPNP), Chiang Mai Province, northern Thailand. Three adult males were collected from a small breeding pond during our survey in July 2020. Based on molecular (two specimens) and morphological (three specimens) evidence, all three specimens from DPHPNP were identified as Tylototriton verrucosus Anderson, 1871, and were characterized by having a blackish background, orange to light orange-brown color markings, segmented vertebral ridge, prominent rib nodules ranging from 13–15 warts, and a rough skin with granules. Hence, this second record of T. verrucosus populations in Thailand confirms a range extension of this species in northern Indochina. Additionally, the morphological variation and conservation status are discussed for the populations from Thailand.

TYLOTOTRITON (TYLOTOTRITON) HOUI HERNANDEZ & DUFRESNES SP. NOV. (FIG. 10) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. o r g: a c t: A D B 2 E B 7 A - E A 1 C - 4 C 9 5 - 9 9 9 6 - EAF78C2FD 96F. Identity and diagnosis: A moderate size Tylototriton slightly flattened in appearance, with an oval head, a prominent vertebral ridge and 11–15 dorsolateral glandular warts. The tail is thick, laterally compressed, and smaller than the SVL. The skin is rough and finely granulated on dorsum and flanks. The background colour is dark, but with extensive orange coloration on the vertebral ridge, the glandular warts, the lower abdomen, the tail, legs as well as large parts of the head. This new species generally resembles T. panaeaensis, T. pulcherrimus and T. Ʋerrucossus, its closest relatives (Fig. 2). It was previously confounded with the shanjing morphotype of T. Ʋerrucosus. Among previous molecular work, T. houi only appears in the fine-scale phylogeography of ‘ T. shanjing ’ by Yu et al. (2013) as the Hengduan endemic clade B, the sister-lineage of clade A (now T. panaeaensis), although this phylogenetic position is not robustly supported (Fig. 1). The new species diverged from the above congeners around the Plio-Pleistocene transition (~2.3 Mya) and features mitochondrial differentiation (0.6–1.0% at 16S, 1.6– 2.4% at Cytb) comparable to other species-level splits in Tylototriton (e.g. T. aeenxianensis / maolanensis and T. pseudoƲerrucosus / taliangensis). Beside molecular characters, T. houi can be distinguished from its close congeners (notably T. pulcherrimus and T. Ʋerrucosus) by its orange abdominal region, an indistinct sagittal crest on the head, distinct nostrils, a broader head and a slimmer body. Moreover, the larvae shows distinctive morphological features compared to T. Ʋerrucosus (shanjing morphotype; Hernandez et al., 2019), such as smaller and darker gills, corrugated and dark tail fins, and a yellow dorsal stripe when reaching larval sizes of 30–36 mm (TTL). Holotype: MZL–46960, adult male found dead on 19 July 2015 by AH, M. Hou, G. Espallargas in Jade Water Village, on the southern slopes of Jade Dragon Snow Mountain (Hengduan massif), Lijiang Prefecture, Yulong Naxi Autonomous County, northern Yunnan Province, China, between 2850 and 3200 m a.s.l. (27.0744°N, 100.1906°E); curated at the Cantonal Museum of Zoology of Lausanne. The specimen was sequenced for 16S and the corresponding lineage is featured in our phylogeny (red circle of the middle clade in Fig. 2). The specimen and the type locality are depicted in Fig. 10. Description of holotype: Poor state due to early decomposition at discovery, which caused discoloration and skin damages. Slim newt (TTL: 87 mm) with fine granulation; laterally compressed tail (TAL: 40 mm), shorter than body (SVL: 47 mm); broad head (HW: 13 mm), larger than body (CW: 11 mm), with distinct nostrils as close to each other’s (IN: 3 mm) than from the eyes (ON: 3 mm); large eyes (ED: 3 mm) laterally disposed; parotids visible; vertebral ridge prominent (discoloured); 16 lateral glandular warts, disposed from axilla to the tail basis; forelimbs (AL: 16 mm) slightly shorter than hind limbs (PL: 18 mm); four fingers and five toes well developed, with no visible webbing; cloaca distinct, with longitudinal vent slit; previously coloured body parts (visible from discoloration patterns) include the lateral sides of the head, the vertebral ridge, the glandular warts, the tail and several underparts (cloaca, hind limbs, forelimbs, gular fold and jaw). Etymology: We coin the new nomen, Tylototriton (Tylototriton) houi, as a tribute to Mian Hou, a leading figure in Chinese herpetology who contributed significant advances in crocodile newt research and taxonomy, including reports on the first known populations of T. (T.) houi (Hernandez et al., 2019). Common names: Hou’s crocodile newt (English), Tylototriton de Hou (French). DiƲersity and distribution: The species is microendemic of the Hengduan Moutains in Yunnan Province, China. It is currently known from only three locations: Lijiang, Peiliang and Shuanghaizi. These populations are genetically similar (Yu et al., 2013; this study). Natural history: The new species inhabits highelevation subalpine meadows surrounded by conifer and mixed broadleaf forests, where permanent or ephemeral ponds (filled by seasonal rains) are available (Hernandez et al., 2019). On average, T. (T.) houi occurs at higher elevations than any other crocodile newt (2714–3200 m a.s.l.). Breeding is aquatic: adults and larvae (depicted in Fig. 10) were observed in vegetated and non-vegetated ponds with roughly neutral pH (6.6–7.7), shared with anuran tadpoles of Bombina maxima (Boulenger, 1905), Rana chaochiaoensis Liu, 1946, Bufo andreaesi Schmidt, 1925, Nanorana yunnanensis (Anderson, 1879). Courtship behaviour has never been observed. Eggs are round, deposited underwater on rocks and submerged terrestrial plants, with capsules of 4.4–5.7 mm in diameter, surrounding embryos of 1.2–1.8 mm (AH pers. obs.). ConserƲation: Although not specifically assessed, T. (T.) houi faces immediate danger of extinction.The few known populations are threatened by anthropogenic pressure related to mass tourism, including water pollution, the introduction of invasive fish (Gambusia Poey, 1854, AH pers. obs.), habitat alteration and destruction, electric fishing. Poaching for traditional Chinese medicine and the pet trade may also represent additional threats.

TYLOTOTRITON (TYLOTOTRITON) HOUI HERNANDEZ & DUFRESNES SP. NOV. (FIG. 10) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. o r g: a c t: A D B 2 E B 7 A - E A 1 C - 4 C 9 5 - 9 9 9 6 - EAF78C2FD 96F. Identity and diagnosis: A moderate size Tylototriton slightly flattened in appearance, with an oval head, a prominent vertebral ridge and 11–15 dorsolateral glandular warts. The tail is thick, laterally compressed, and smaller than the SVL. The skin is rough and finely granulated on dorsum and flanks. The background colour is dark, but with extensive orange coloration on the vertebral ridge, the glandular warts, the lower abdomen, the tail, legs as well as large parts of the head. This new species generally resembles T. panaeaensis, T. pulcherrimus and T. Ʋerrucossus, its closest relatives (Fig. 2). It was previously confounded with the shanjing morphotype of T. Ʋerrucosus. Among previous molecular work, T. houi only appears in the fine-scale phylogeography of ‘ T. shanjing ’ by Yu et al. (2013) as the Hengduan endemic clade B, the sister-lineage of clade A (now T. panaeaensis), although this phylogenetic position is not robustly supported (Fig. 1). The new species diverged from the above congeners around the Plio-Pleistocene transition (~2.3 Mya) and features mitochondrial differentiation (0.6–1.0% at 16S, 1.6– 2.4% at Cytb) comparable to other species-level splits in Tylototriton (e.g. T. aeenxianensis / maolanensis and T. pseudoƲerrucosus / taliangensis). Beside molecular characters, T. houi can be distinguished from its close congeners (notably T. pulcherrimus and T. Ʋerrucosus) by its orange abdominal region, an indistinct sagittal crest on the head, distinct nostrils, a broader head and a slimmer body. Moreover, the larvae shows distinctive morphological features compared to T. Ʋerrucosus (shanjing morphotype; Hernandez et al., 2019), such as smaller and darker gills, corrugated and dark tail fins, and a yellow dorsal stripe when reaching larval sizes of 30–36 mm (TTL). Holotype: MZL–46960, adult male found dead on 19 July 2015 by AH, M. Hou, G. Espallargas in Jade Water Village, on the southern slopes of Jade Dragon Snow Mountain (Hengduan massif), Lijiang Prefecture, Yulong Naxi Autonomous County, northern Yunnan Province, China, between 2850 and 3200 m a.s.l. (27.0744°N, 100.1906°E); curated at the Cantonal Museum of Zoology of Lausanne. The specimen was sequenced for 16S and the corresponding lineage is featured in our phylogeny (red circle of the middle clade in Fig. 2). The specimen and the type locality are depicted in Fig. 10. Description of holotype: Poor state due to early decomposition at discovery, which caused discoloration and skin damages. Slim newt (TTL: 87 mm) with fine granulation; laterally compressed tail (TAL: 40 mm), shorter than body (SVL: 47 mm); broad head (HW: 13 mm), larger than body (CW: 11 mm), with distinct nostrils as close to each other’s (IN: 3 mm) than from the eyes (ON: 3 mm); large eyes (ED: 3 mm) laterally disposed; parotids visible; vertebral ridge prominent (discoloured); 16 lateral glandular warts, disposed from axilla to the tail basis; forelimbs (AL: 16 mm) slightly shorter than hind limbs (PL: 18 mm); four fingers and five toes well developed, with no visible webbing; cloaca distinct, with longitudinal vent slit; previously coloured body parts (visible from discoloration patterns) include the lateral sides of the head, the vertebral ridge, the glandular warts, the tail and several underparts (cloaca, hind limbs, forelimbs, gular fold and jaw). Etymology: We coin the new nomen, Tylototriton (Tylototriton) houi, as a tribute to Mian Hou, a leading figure in Chinese herpetology who contributed significant advances in crocodile newt research and taxonomy, including reports on the first known populations of T. (T.) houi (Hernandez et al., 2019). Common names: Hou’s crocodile newt (English), Tylototriton de Hou (French). DiƲersity and distribution: The species is microendemic of the Hengduan Moutains in Yunnan Province, China. It is currently known from only three locations: Lijiang, Peiliang and Shuanghaizi. These populations are genetically similar (Yu et al., 2013; this study). Natural history: The new species inhabits highelevation subalpine meadows surrounded by conifer and mixed broadleaf forests, where permanent or ephemeral ponds (filled by seasonal rains) are available (Hernandez et al., 2019). On average, T. (T.) houi occurs at higher elevations than any other crocodile newt (2714–3200 m a.s.l.). Breeding is aquatic: adults and larvae (depicted in Fig. 10) were observed in vegetated and non-vegetated ponds with roughly neutral pH (6.6–7.7), shared with anuran tadpoles of Bombina maxima (Boulenger, 1905), Rana chaochiaoensis Liu, 1946, Bufo andreaesi Schmidt, 1925, Nanorana yunnanensis (Anderson, 1879). Courtship behaviour has never been observed. Eggs are round, deposited underwater on rocks and submerged terrestrial plants, with capsules of 4.4–5.7 mm in diameter, surrounding embryos of 1.2–1.8 mm (AH pers. obs.). ConserƲation: Although not specifically assessed, T. (T.) houi faces immediate danger of extinction.The few known populations are threatened by anthropogenic pressure related to mass tourism, including water pollution, the introduction of invasive fish (Gambusia Poey, 1854, AH pers. obs.), habitat alteration and destruction, electric fishing. Poaching for traditional Chinese medicine and the pet trade may also represent additional threats., Published as part of Dufresnes, Christophe & Hernandez, Axel, 2023, Towards completing the crocodile newts' puzzle with all-inclusive phylogeographic resources, pp. 620-640 in Zoological Journal of the Linnean Society 197 on pages 636-637, DOI: 10.1093/zoolinnean/zlac038, http://zenodo.org/record/7695335, {"references":["Yu G, Zhang M, Rao D, Yang J. 2013. Effect of Pleistocene climatic oscillations on the phylogeography and demography of red knobby newt (Tylototriton shanjing) from southwestern China. PLoS One 8: e 56066.","Hernandez A, Hou M, Espallargas G. 2019. On the northernmost populations of Tylototriton shanjing Nussbaum, Brodie & Yang, 1995 in Yunnan province, China. Bulletin de la Societe Herpetologique de France 169: 3 - 10."]}

The distribution of the Ziegler's Crocodile Newt Tylototriton ziegleri Nishikawa, Matsui & Nguyen, 2013 in China has been controversial. This species was originally recorded uncertainly from Guangxi Autonomous Region, China. Subsequently, this species was recorded from Yunnan and Guangdong provinces, China. Thereafter, the record from Guangdong was denied and the record from Yunnan was questioned.Two specimens of Tylototriton Anderson, 1871 were collected from Wenshan Prefecture, Yunnan Province, China, in 2021. Phylogenetically, the sequences of these two specimens clustered with the sequences of T. ziegleri (including the holotype) from its type locality with strong support, and morphologically agree well with the original description of T. ziegleri. We confirm the record of T. ziegleri in China and present detailed collection site and morphological description of the specimens from China. In addition, we found that T. panwaensis Grismer, Wood, Quah, Thura, Espinoza & Murdoch, 2019 may be the synonym of T. verrucosus Anderson, 1871. We discussed the relationship between T. verrucosus and T. panwaensis.

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).

The Tylototriton asperrimus complex from northern Vietnam is reviewed based on morphological comparisons and analysis of the mitochondrial marker NADH dehydrogenase subunit 2 (ND2). Based on molecular divergences, which were revealed to be higher than in other congeners, in concert with morphological differences, two new species and one subspecies are described herein: Tylototriton pasmansisp. nov. differs from T. asperrimussensu stricto by 3.2 to 3.6 % genetic divergence and a combination of distinct morphological characters, such as head slightly longer than wide, distinct mid-dorsal ridge, relatively wide distance between the eyes, tips of fingers reaching the eye when foreleg is laid forward, labial and gular folds present, central belly skin with tubercles shaped like transverse wrinkles and distinct, pointy to round rib nodules. The population of T. pasmansisp. nov. consists of two subclades, the nominotypic one occurring on the eastern side of the Da River (or Black River, including Hoa Binh and Phu Tho provinces), and another occurring on the western side (including Son La and Thanh Hoa provinces). These two subclades differ by 2.5 to 3.1 % genetic divergence and distinct morphological characters. The western subclade is herein described as Tylototriton pasmansi obstissp. nov., which differs from the nominotypic form by a wider head, longer and narrower snout, shorter femur length, and an overall less granulose skin, without an increased concentration of warts on the body sides. A second new species, Tylototriton sparreboomisp. nov. is described from Lai Chau Province. It differs from T. asperrimussensu stricto by 4.1 to 4.2 % and from Tylototriton pasmansisp. nov. by 3.6 to 4.5 % genetic divergences as well as by a combination of distinct morphological characters, such as head longer than wide, tips of fingers reaching nostril when foreleg adpressed along head, rib nodules distinct, round and relatively enlarged, and wide distance between the eyes.

Morphological observations (LM, TEM, SEM) of olfactory organs were conducted on three representatives of the family Salamandridae which differ in post-metamorphic habitat: the terrestrial fire salamander Salamandra salamandra, the semiaquatic alpine newt Ichthyosaura alpestris and the semiaquatic Himalayan newt Tylototriton verrucosus which exhibit increasing specialization towards life in water. We demonstrated variability in proportional size and shape of the main olfactory chamber (MOC), as well as in the degree of development of the vomeronasal organ (VNO) among the three species. In the fire salamander, the olfactory sensory epithelium was distributed throughout the MOC, while in the alpine and Himalayan newt the olfactory sensory epithelium was organized in strips. The presence of two types of olfactory receptor neurons (ciliated receptor neurons, microvillar receptor neurons) was confirmed in olfactory organs of all examined species. For the first time in amphibians, a subtype of ciliated receptor neuron – the giant neuron (rod-shape neuron), was detected in the MOC of all three salamandrids. The olfactory epithelium also consists of ciliated supporting cells and secretory supporting cells. The presence of goblet cells, mucociliary epithelial cells and squamous epithelial cells was demonstrated in the olfactory organs of all examined species. Our results indicate that the olfactory organs of salamandrids may differ in size, shape and distribution of olfactory epithelium depending on habitat type during the post-metamorphic stages of ontogeny.

A new species of crocodile newt from populations on opposite sides of the Ayeyarwady Basin, Myanmar, previously referred to as Tylototriton cf. verrucosus, is described on the basis of colour patte...

We describe a new species of the genus Tylototriton from Ingyin Taung Mt., Mohnyin Township, Kachin State, Myanmar, based on morphological and molecular evidence. The new species is assigned to the subgenus Tylototriton s. str. and is clearly distinct from all known congeners by the following characters: medium body size; thin, long tail, lacking lateral grooves; rough skin; truncate snout; wide, protruding supratemporal bony ridges on head, beginning at anterior corner of orbit; weak, almost indistinct sagittal ridge; long, thin limbs, broadly overlapping when adpressed along body; distinct, wide, non-segmented vertebral ridge; 13 or 14 rib nodules; brown to dark-brown background coloration with dull orange-brown to yellowish-brown markings on labial regions, parotoids, rib nodules, whole limbs, vent, and ventral tail ridge. We also briefly discuss biogeography and species diversity of the genus Tylototriton in Myanmar.

Niche conservatism underlies the process of speciation in amphibians on the global scale. However, high tropical diversity is partly attributable to niche shifts within evolutionary lineages. Crocodile newts (genera Echinotriton and Tylototriton) are a monophyletic group endemic to south-east Asia, and are one of the few lineages of urodeles that appear in tropical regions. Here, we examined the patterns of ecological diversification within the crocodile newts based on 207 records of 23 species. The niche space was described according to topographic, climatic variables (temperature, rainfall, and seasonality) and forest cover. Our analysis showed that the occurrence of most of these species was strongly associated with the precipitation regime and seasonality. The diversification patterns of the crocodile newts are complex, showing both conservatism and divergence. These patterns can be related to the Tibetan uplift phase during the Neogene, resulting in the emergence of new montane habitats and subsequent changes in the palaeo-drainage system and climate. The range of ecological diversification within a lineage is possibly the result of several interacting processes, including environmental heterogeneity, the regional climatic history, and physiological constraints on species.

Tylototriton ziegleri is distributed in northern Vietnam and is a new member of the genus Tylototriton. The conservation status of this species has not yet been assessed by the IUCN, but several other species of Tylototriton have been listed as near threatened or vulnerable due to habitat destruction and illegal hunting. We determined the complete sequence and structural characteristics of the mitochondrial genome of T. ziegleri. The entire mitogenome is 16,266 bp in length, and contains 13 protein-coding genes, 22 tRNA genes, two rRNA genes, a control region, and a non-coding region. The length, gene arrangement, and composition are similar to mitogenomes of other species of Tylototriton. A phylogenetic tree based on the mitogenome sequences of several Asian species within Salamandridae confirmed the genus Tylototriton as monophyletic, with T. ziegleri being the sister species of T. asperrimus and both constituting the sister group of T. wenxianensis. This work provides basic molecular data which are critical for further genetic research and conservation of the genus Tylototriton.

The diversity of the vertebrate cranial shape of phylogenetically related taxa allows conclusions on ecology and life history. As pleurodeline newts (the genera Echinotriton, Pleurodeles and Tylototriton) have polymorphic reproductive modes, they are highly suitable for following cranial shape evolution in relation to reproduction and environment. We investigated interspecific differences externally and differences in the cranial shape of pleurodeline newts via two‐dimensional geometric morphometrics. Our analyses also included the closely related but extinct genus Chelotriton to better follow the evolutionary history of cranial shape. Pleurodeles was morphologically distinct in relation to other phylogenetically basal salamanders. The subgenera within Tylototriton (Tylototriton and Yaotriton) were well separated in morphospace, whereas Echinotriton resembled the subgenus Yaotriton more than Tylototriton. Oviposition site choice correlated with phylogeny and morphology. Only the mating mode, with a random distribution along the phylogenetic tree, separated crocodile newts into two morphologically distinct groups. Extinct Chelotriton likely represented several species and were morphologically and ecologically more similar to Echinotriton and Yaotriton than to Tylototriton subgenera. Our data also provide the first comprehensive morphological support for the molecular phylogeny of pleurodeline newts., We applied geometric morphometrics to cranial morphology in extant and extinct crocodile newts. Our data provides the first comprehensive morphological support for the molecular phylogeny of these newts. Further, we could show that life history traits correlate with head shape and that fossil Chelotriton show similarities in shape with some extant taxa allowing conclusions on its ecology.

The existence of two or more distinctly coloured phenotypes among individuals of an interbreeding population is known as colour polymorphism. In amphibians, this phenomenon is pervasive among anurans, but rare or absent among salamanders and caecilians, respectively. Here, we examine whether various distinct phenotypes of Salamandra salamandra in North Spain, used as a basis to describe the subspecies S. s. bernardezi and S. s. alfredschmidti, indeed warrant separate taxonomic status or that these co-occur and belong to a single taxon. Based on a sample of 1147 individuals from 27 local populations, six phenotype classes were designated. Although two phenotypes that are attributable to S. s. alfredschmidti show some degree of geographical restriction, these co-occur with those representing typical S. s. bernardezi. A fifth phenotype class could not be unambiguously attributed to either subspecies due to an overlap in previously suggested diagnostic characteristics. Mitochondrial (cytochrome b) and nuclear (β-fibrinogen) DNA analyses revealed S. s. alfredschmidti to be nested within several subclades of S. s. bernardezi, without displaying unique lineages. Furthermore, no significant divergence was recovered by means of niche overlap analyses. As a result, we revoke the subspecies status of S. s. alfredschmidti, which should be regarded as a junior synonym of S. s. bernardezi. The current findings confirm the existence of colour polymorphism in S. salamandra and the family Salamandridae, which provides exciting possibilities for future research.

Tylototriton shanorum, an endemic species of Myanmar, is threatened by illegal capture and environmental destruction. Here, the complete mitochondrial genome (mitogenome) of T. shanorum is described. The entire mitogenome sequence of T. shanorum is 17,096 bp long, containing 13 protein coding genes, two ribosomal RNA, 22 transfer RNA genes, one control region. In addition, two additional 3′ Cytb like sequences and two complete tRNAThr genes, one pseudo tRNAThr, and three noncoding sequences (NC1, NC1′ and NC2) were situated between Cytb and tRNAPro genes. The new complete mitogenome sequence of T. shanorum will be useful for conservation genetics, evolutionary biology, phylogeography, as well as phylogenetic relationships studies of this threatened species.

A phylogenetic taxonomic analysis of a monophyletic subgroup of the salamandrid genus Tylototriton revealed that a newly discovered population from Ngar Su Village, 1 km south of Ywangan, Shan State, Myanmar is a new species and most closely related to T. shanorum from approximately 80 km to the west in the vicinity of Taunggyi, Shan State. Tylototriton ngarsuensis sp. nov. differs from other closely related species of Tylototriton on basis of varying combinations of characteristics associated with it shorter head, larger size, rib nodule morphology, and overall drab, very dark, coloration, along with its genetic differentiation. Tylototriton ngarsuensis sp. nov. also appears to breed later in the year than most other species. Unfortunately, this species like many other Asian newts, is being harvested for the pet and medicinal trade and given its restricted distribution, this could pose a serious threat to its long-term survival.

The natural history of newly discovered species among the genus Tylototriton in South-Eastern Asia is poorly known to date, and most of the species are threatened by extinction. The description of their ecology is crucial to determine their survival status and successful reproduction for conservation purposes. In this study, we assessed the microhabitat preferences of Tylototriton yangi at the type locality, Gejiu, Yunnan province, China, with the Visual Encounter Survey method and by using new important materials. The main habitat is located within a humid subtropical climate and is composed of a large karstic mountain containing some patch of secondary mixed forests and plantations such as tomato fields, pepper and cabbage that are surrounded by scrubs, grasses and small ponds. We provided a complete diagnosis with new morphological data analysing ten adult individuals from the type series to establish a formal description which is absent from the first original description. We compared our results with other related taxa of the genus Tylototriton to updated key determinants between similar species. Thus, we analysed the population structure and the behaviour of T. yangi during a period of ten years, 2008–2017, to assess its survival status and to determine the main threat factors. A total of 186 specimens were found during the whole study predicting a small population inhabiting the type locality, Gejiu. Consequently, T. yangi is now regarded as a highly threatened species that need more attention of Chinese authorities and we suggest regarding it as Endangered due to its small distribution range, the low number of adult specimens observed and ongoing exploitation.

In vertebrates, the relative proportion of the number of trunk and caudal vertebrae is an important determinant of body shape. While among amphibians frogs and toads show low variation in vertebrae numbers, in salamanders the numbers of trunk and caudal vertebrae vary widely, giving rise to phenotypes in the range from short-bodied and long-tailed to long-bodied and short-tailed. We analysed vertebral numbers in the family Salamandridae in a phylogenetic context and calculated the relationship between vertebral changes and changes in climate and other environmental parameters. A significant association was found between morphological change with precipitation and temperature. However, annual precipitation affected the two main groups of salamandrid salamanders differently, with trunk elongation in the terrestrial ‘true salamanders’ and tail elongation in the more aquatic ‘newts’. A - male biased - sexual dimorphism was only observed in Lissotriton vulgaris vulgaris in the number of trunk vertebrae and in Ommatotriton ophryticus and Lissotriton species for the number of caudal vertebrae. Our data indicated that the number of trunk and caudal vertebrae are highly evolvable traits with frequent evolutionary reversals. In some groups (e.g. Cynops, Lyciasalamandra, Neurergus and the Laotriton- Pachytriton-Paramesotriton clade) the number of trunk vertebrae is stable, while in many groups it is subject to change (e.g. Tylototriton). This latter, species-rich genus appears to be an excellent group to further test effects of the environment on body shape.

We assessed taxonomic relationship of Tylototriton daweishanensis Zhao, Rao, Liu, Li and Yuan, 2012 and T. yangi Hou, Li and Lu, 2012 using mitochondrial DNA sequence data and found them to be as closely related as to be regarded as conspecific. This result, together with available morphological information, strongly indicates that T. daweishanensis is a junior synonym of T. yangi.

We describe a new species of Tylototriton from northwestern Vietnam and northern Thailand based on morphological and molecular evidence. Tylototriton anguliceps sp. nov. is distinguishable from all the other congeners by the bright to dark orange markings on the head, body, and tail, prominent dorsal and dorsolateral ridges (crests) on the head, skeletal connection between maxillary and pterygoid, and unique mitochondrial and nuclear DNA sequences. Our molecular data show that the new species is nested within the clade comprising T. uyenoi, T. shanjing, T. verrucosus, and T. yangi. The new species is expected to be recorded from other countries in the Indochina region such as southern China, western Myanmar and northern Laos in the future.

Abstarct Tylototriton anhuiensis (Salamandridae, Urodela), collected from Yaoluoping Nature Reserve, was identified as a new species. The complete mitogenome sequence of T. anhuiensis is 16,259 by in length, including 13 protein-coding genes, 22 tRNA genes, 2 rRNA genes, and D-loop region. The base composition of the mitogenome was 33.6%A, 26.3% C, 14.5% G, and 25.6% T. The ND6 subunit gene and eight tRNA genes were encoded on the L-strand, the others were encoded on the H-strand.

A new species of salamandrid is named and described from the Miocene of Randeck Maar, a long-term vol- canic lake deposit in southern Germany. Based on a fully articulated specimen, Ichthyosaura randeckensis is named and described as a 3-4 cm long newt with a robust posterolateral process of the frontal, a heavily ornamented skull roof, and a premaxilla with spike-like, parallel alary processes framing a slender fontanelle posteriorly. In a phylogenetic analysis of 14 salamandrid taxa and 38 morphological characters, I. randeckensis is found to nest with I. alpestris, with which it shares apomorphic character states in the morphology of premaxilla and nasal. The present analysis employed more morpholog- ical characters than hitherto considered, highlighting the need to conduct more osteological studies of extant salamanders. Its results are largely consistent with recent molecular phylogenetic studies: Tylototriton and Pleurodeles form a clade, with Notophthalmus and Euproctus falling within unresolved trichotomies, followed by Cynops and Ommatotriton as successive crownward clades, then Triturus sensu stricto (T. marmoratus and T. cristatus), and finally Lissotriton and Ichthyosaura as terminal sister taxa. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Crocodile newts, which constitute the genera Echinotriton and Tylototriton, are known as living fossils, and these genera comprise many endangered species. To id entify mitochondrial (mt) genes suitable for future population genetic analyses for endangered taxa, we determined the complete nucleotide sequences of the mt genomes of the Japanese crocodile newt Echinotriton andersoni and Himalayan crocodile newt Tylototriton verrucosus. Although the control region (CR) is known as the most variable mtDNA region in many animal taxa, the CRs of crocodile newts are highly conservative. Rather, the genes of NADH dehydrogenase subunits and ATPase subunit 6 were found to have high sequence divergences and to be usable for population genetics studies. To estimate the inter-population divergence ages of E. andersoni ende mic to the Ryukyu Islands, we performed molecular dating analysis using whole and partial mt genomic data. The estimated di vergence ages of the inter-island individuals are older than the paleogeographic segmentation ages of the islands, suggesting that the lineage splits of E. andersoni populations were not caused by vicariant events. Our phylogenetic analysis with pa rtial mt sequence data also suggests the existence of at least two more undescribed species in the genus Ty lototriton. We also found unusual repeat s equences containing the 3′ region of cytochrome apoenzyme b gene, whole tRNAThr gene, and a noncoding region (the T-P noncoding region characteristic in caudate mtDNAs) from T. v errucosus mtDNA. Similar repeat sequences were found in t wo other Tylototriton species. The Ty lototriton taxa with the repeats become a monophyletic group, indicating a single origin of the repeat sequences. The intraa nd inter-specific comparisons of the repeat sequences suggest the occurrences of homologous recombination-based concerted evolution among the repeat sequences.

Tylototriton podichthys sp. nov. (Fig. 6) Tylototriton shanjing Matsui 2013: 187; Nishikawa, Khonsue, Pomchote & Matsui 2013 a: 264 (part); Sparreboom 2014: 370 (part). Tylototriton verrucosus verrucosus (part) Le, Nguyen, Nishikawa, Nguyen, Pham, Matsui, Bernardes & Nguyen 2015: 39. Holotype: NCSM 77721 (field tag BLS 12909), adult female, Laos, Luang Phabang Province, Phoukhoun District, Nam Madao Village, 19 º 18 ’ 0.3 ”N 102 º 34 ’ 25.2 ”E, 1,189 m elev., coll. 17 August 2009 by Somphouthone Phimmachak. Paratypes: NCSM 77722 – 23 (two females), NCSM 77724 – 26 (three males), same data as holotype. NUOL 0 0 393 (one male), same data as holotype except 19 º 16 ’ 59.9 ”N 102 º 34 ’ 41.6 ”E, 1,287 m elev., coll. 8 June 2009. NUOL 00395– 97 (three males), same data as holotype except 19 º 18 ’06.8”N 102 º 34 ’ 24.8 ”E, 1,287 m elev., 8–10 June 2009. NCSM 82935 (one female), same data as holotype except 19.30189 ºN 102.57356 ºE, 1,287 m elev., coll. 10 July 2012. NCSM 82939 – 40, NUOL 00402–03, NUOL 0 0 411 (five females), NCSM 82941 – 43, NUOL 00409– 10, NUOL 0 0 438, (six males), Laos, Xieng Khouang Province, Kham District, Yord Lieng Village, Mor Stream, 19.83986 ºN 103.70425 ºE, 1,481 m elev., coll. 13 June 2011 by Somphouthone Phimmachak. NUOL 0 0 436, NUOL 0 0 445 (two females), NUOL 0 0 435, NUOL 0 0 444 (two males), Laos, Xieng Khouang Province, Kham District, Yord Lieng Village, Mor Stream, 19.83972 ºN 103.70259 ºE, 1,493 m elev., coll. 6 July 2012 by Somphouthone Phimmachak. NCSM 82936, NUOL 0 0 412 (two females), Laos, Huaphanh Province, Viengthong District, Phou Louey National Protected Area, Sakok Village, headwaters of Nam Long Stream, 20.22556 ºN 103.23122 ºE, 1,348 m elev., coll. 16–17 June 2011 by Somphouthone Phimmachak. NCSM 82937 – 38 (two males), Laos, Huaphanh Province, Viengthong District, Phou Louey National Protected Area, Sakok Village, headwaters of Nam Long Stream, 21.22669 ºN 104.23606 ºE, 1,303 m elev., coll. 18 June 2011 by Somphouthone Phimmachak. NUOL 0 0 413 (one male), Laos, Huaphanh Province, Viengthong District, Phou Louey National Protected Area, Sakok Village, headwaters of Nam Long Stream, 21.23153 ºN 104.23050 ºE, 1,317 m elev., coll. 17 June 2011 by Somphouthone Phimmachak. Referred material. NCSM 82933 (22 larvae), same data as NUOL 0 0 435 except 22 August 2012. NCSM 82934 (nine larvae), NUOL 0 0 450 (larvae, number of individuals not recorded), same data as NCSM 82939 except coll. 13–16 October 2010 by Somphouthone Phimmachak and Sengvilay Seateun. NUOL 00406–08, NUOL 0 0 414 (four metamorphs), same data as NUOL 0 0 435 except coll. 10 December 2012. NUOL 0 0 394 (one eft), same data as NUOL 0 0 395. NCSM 82944 – 46, NUOL 00439– 42 (seven efts), same data as NCSM 82939. NCSM 82948 – 51 (four efts), same data as NUOL 0 0 435 except coll. 13–16 December 2012. NCSM 82947 (one eft), same data as NCSM 82937. NUOL 0 0 446 (one eft), same data as NUOL 0 0 413. Subgenus: The new species is assigned to the subgenus Tylototriton Dubois & Raffaëlli 2009 (= T. verrucosus species group of Fei et al. 2005) based on its phylogenetic position (Fig. 1) and having orange coloration on limbs and lateral surfaces of tail (Nishikawa et al. 2014). Etymology. The specific epithet is a noun in apposition taken from pod Gr., foot, and ichthys Gr., fish, in reference to the Lao language name pa theen, meaning “fish with feet,” that is widely used for newts (Tylototriton and Laotriton) in Laos. Diagnosis. Tylototriton podichthys sp. nov. is a Tylototriton (subgenus Tylototriton) having the combination of an indistinct glandular ridge on midline of crown; distinct, rounded rib nodules with diameter equivalent to or greater than that of eye; parotoid oriented parallel to body axis in lateral view; thick, glandular, vertebral tubercular ridge; rough, glandular skin on cranial crest; uniformly orange cranial crest and parotoids; orange markings separated between rib nodules; dark coloration on ventral surfaces of limbs; and dark coloration on digit tips. Description of holotype. Habitus stout. Head narrower than body, longer than wide, slightly sloping in profile. Snout short, rounded in dorsal view, slightly rounded in profile, exceeding beyond lower jaw. Nostrils close to snout tip, visible from above. Vomeropalatine tooth series in an inverted, narrow V-shape, anteriorly in contact and reaching choanae. Tongue oval, attached at anterior floor of mouth. Gular fold present. Indistinct glandular ridge on midline of crown from level of center of eye to level of corner of mouth. Strong glandular cranial crest on outer margin of crown from level of nostril, weakly curved medially at posterior end, continuous with parotoid. Parotoids enlarged, projecting posteriorly, oriented parallel to body axis in lateral view. Strong vertebral tubercular ridge from posterior end of crown to dorsal margin of tail, visible on anterior one-third of tail, weakly separated from ridge on midline of crown. Dorsolateral row of large glandular warts (rib nodules) on each side from level of axilla to anterior one-third of tail, 16 on left and 15 on right between axilla and level of posterior margin of vent, distinctly knob-like anteriorly, becoming smaller posteriorly, third on both sides the largest. Smaller, glandular warts on most of remaining dorsal and ventral surfaces, warts on crown, nape, and ventrolateral region with clusters of glands, those on throat, tail, and ventral surfaces of limbs granular, those on belly arranged in striations perpendicular to body axis, with a smooth, glandular, ovoid patch of skin on chest. Four fingers, five toes, all without webbing. Tail laterally compressed, tapering posteriorly, narrow dorsal fin, smooth ventral ridge, tip rounded in profile. Color of holotype in life. Dorsal surface of body, nape, and lores black. Throat, chest, belly, and ventral surfaces of limbs black with limited, irregular orange mottling. Remainder of head, parotoids, vertebral ridge, rib nodules, ventral margin of lower jaw, dorsal surfaces of limbs, ventral surfaces of hands and feet, cloaca, and tail orange, with brightest orange on parotoids, vertebral ridge and rib nodules. Tips of all digits dark brown. Iris brown, pupil black. Color of holotype in preservative. Orange faded to beige or brown. Variation. Some paratypes (e.g., females NCSM 82939 – 40, male NCSM 82943) have much darker orange coloration in life and preservative than holotype (Fig. 6 C). Some paratypes have dorsolateral rows of rib nodules extending only to base of tail. Some paratypes have truncate snout in dorsal view. Females are heavier, have a more robust body, longer body (SVLA, SVLP, TTL), and shorter vent slits than males (Phimmachak et al., in press). Tail tip acuminate rather than rounded in profile in some paratypes. Measurements of adults are summarized in Table 7. Distribution and natural history. Tylototriton podichthys sp. nov. is currently known only from Luang Phabang, Xieng Khouang and Huaphanh Provinces, Laos, from 1,189–1,493 m elevation (Fig. 3). A reported locality of Phu Pan, Xam Neua (Matsui 2013; Nishikawa et al. 2013 a; Le et al. 2015) is also in Huaphanh Province. A reported locality of Nam Lieng (Sparreboom 2014) is the same as the paratype locality in Kham District, Xieng Khouang Province, Laos. Sparreboom (2014) provided a photograph of habitat in Xieng Khouang Province. The natural history and general ecology of the new species is extensively reviewed in Phimmachak et al. (in press). Comparisons. Tylototriton podichthys sp. nov. differs from all species in the T. asperrimus group (subgenus Yaotriton) by having orange coloration on limbs and lateral surfaces of tail. The new species is less readily differentiated from some co-members of the T. verrucosus group (subgenus Tylototriton; Figs. 7–9). Tylototriton podichthys sp. nov. differs from T. anguliceps by having an indistinct glandular ridge on midline of crown (distinct ridge on midline of crown in anguliceps) and by having more robust males (males distinctly slimmer in anguliceps). Tylototriton podichthys sp. nov. differs from T. kweichowensis and T. pseudoverrucosus by having separated orange markings on rib nodules (orange markings connected, forming continuous dorsolateral lines from parotoid to tail in kweichowensis and pseudoverrucosus). Tylototriton podichthys sp. nov. differs from T. shanjing by having an indistinct glandular ridge on midline of crown (distinct ridge on midline of crown in shanjing), no sharp contrast between orange crown and black nape (sharp contrast in shanjing), dark coloration on ventral surfaces of limbs (orange coloration on ventral surfaces of limbs in shanjing), and dark digits tips (uniformly orange digit tips in shanjing). Tylototriton podichthys sp. nov. differs from T. shanorum by having large, rounded rib nodules with diameter equivalent to or greater than that of eye (small, slightly elongated rib nodules with diameter less than that of eye in shanorum), and by having a thick, glandular, vertebral tubercular ridge (distinctly narrower and less glandular in shanorum). Tylototriton podichthys sp. nov. differs from T. taliangensis by having rib nodules (absent in taliangensis) and orange coloration on body (orange coloration limited to parotoid, digit tips, vent, and ventral surface of tail in taliangensis). Tylototriton podichthys sp. nov. differs from T. uyenoi by having males with shorter heads (HL 16.9–21.3, n = 18, in podichthys; HL 19.1– 25.5, n = 7, in uyenoi; Wilcoxon test, Z=- 2.058, pn = 18, in podichthys; HW 16.0– 19.6, n = 7, in uyenoi; Wilcoxon test, Z=- 3.571, puyenoi), and having the parotoid oriented parallel to body axis in lateral view (parotoid oriented obliquely downward relative to body axis in lateral view in uyenoi). Tylototriton podichthys sp. nov. differs from T. verrucosus by having distinctly contrasting coloration between body and the cranial crest, parotoids, vertebral ridge and rib nodules (indistinct contrasting coloration in verrucosus), by having an indistinct glandular ridge on midline of crown (distinct ridge on midline of crown in verrucosus), rough, glandular skin on cranial crest (smooth in verrucosus), and parotoid oriented parallel to body axis in lateral view (parotoid oriented obliquely downward relative to body axis in lateral view in verrucosus). Tylototriton podichthys sp. nov. differs from T. yangi by having uniformly orange cranial crest and parotoids (orange coloration restricted to posterior margin of cranial ridge and parotoids in yangi).

Tylototriton anguliceps Le, Nguyen, Nishikawa, Nguyen, Pham, Matsui, Bernardes & Nguyen 2015 Tylototriton shanjing Group 1 Nishikawa, Khonsue, Pomchote & Matsui 2013 a: 262. Referred material. NCSM 82952 ��� 53, NUOL 0 0 420 (three males), NUOL 0 0 419 (one female), NUOL 0 0 448 (one eft), Laos, Luang Namtha Province, Viengphoukha District, Phou Ya Khao Village, Nam Long Stream, 20.89236 ��N 101.08519 ��E, 1,466 m elev., coll. 8 July 2011 by Somphouthone Phimmachak and Sengvilay Seateun (Figs. 4 C���H). Measurements are summarized in Table 5. Distribution and natural history. Specimens were collected during the day under grass in shrub forest 2���6 m from a 5���7 m wide stream. The shrub forest had been burned within the previous five years. This is the first report of the species in Laos, although the type locality at Muong Nhe, Dien Bien Province, Vietnam is very near to the border of Laos, and so it was expected there (Le et al. 2015). Elsewhere the species is known from Thuan Chau, Son Lao Province, Vietnam, and Doi Lahnga, Chiang Rai Province, Thailand (Le et al. 2015; Fig. 2)., Published as part of Phimmachak, Somphouthone, Aowphol, Anchalee & Stuart, Bryan L., 2015, Morphological and molecular variation in Tylototriton (Caudata: Salamandridae) in Laos, with description of a new species, pp. 285-310 in Zootaxa 4006 (2) on page 297, DOI: 10.11646/zootaxa.4006.2.3, http://zenodo.org/record/237189, {"references":["Le, D. T., Nguyen, T. T., Nishikawa, K., Nguyen, S. L. H., Pham, A. V., Matsui, M., Bernardes, M. & Nguyen, T. Q. (2015) A new species of Tylototriton Anderson, 1871 (Amphibia: Salamandridae) from northern Indochina. Current Herpetology, 34, 38 - 50. http: // dx. doi. org / 10.5358 / hsj. 34.38","Nishikawa, K., Khonsue, W., Pomchote, P. & Matsui, M. (2013 a) Two new species of Tylototriton from Thailand (Amphibia: Urodela: Salamandridae). Zootaxa, 3737 (3), 261 - 279. http: // dx. doi. org / 10.11646 / zootaxa. 3737.3.5"]}

Tylototriton notialis Stuart, Phimmachak, Sivongxay & Robichaud 2010 Tylototriton asperrimus (part) Yuan, Jiang, Lü, Yang, Nguyen, Nguyen, Jin & Che 2011: 583. Tylototriton notialis Nishikawa, Matsui & Nguyen 2013 b: 36. Referred material. FMNH 271120 (male paratype), FMNH 271121 (male holotype), FMNH 271125 (one larva), FMNH 271129 (nine larvae), Laos, Khammouan Province, Boualapha District, Nakai-Nam Theun National Protected Area, Nam On River catchment, Phou Ak escarpment, 17 º 38 ’ 39.6 ”N 105 º 44 ’ 12.3 ”E, 980 m elev., coll. 26–29 May 2007 by Bryan L. Stuart, Somphouthone Phimmachak and Niane Sivongxay. FMNH 271122 (female paratype), same data as holotype except 17 º 39 ’03.4”N 105 º 44 ’ 25.2 ”E, ca. 1,000 m elev., coll. 22 May 2006 by William G. Robichaud. NUOL 0 0 443 (one topotype female), same data as types except 17.64514 ºN 105.73681 ºE, 984 m elev., coll. 14 May 2013 by Somphouthone Phimmachak, Bryan L. Stuart, and Jennifer A. Sheridan (Figs. 4 A–B). Measurements are summarized in Table 4. Distribution and natural history. This is the second report of the species in Laos, but from the same (and only) known locality in Laos. The types were collected day and night in semi-evergreen mixed with pine forest, on a stream bottom or the forest floor (Stuart et al. 2010). The larvae were found at night on the bottom of a 3 m wide stream with slow current, sand and bedrock substrate, and leaf litter-filled potholes (Stuart et al. 2010). The newly reported NUOL topotype was taken at 1115 h after heavy rain under leaf litter and pine needles at the base of a 0.3 m diameter-at-breast-height (DBH) pine tree, 4 m from a stream pool, in semi-evergreen mixed with pine forest. Elsewhere, the species has been reported from adjacent Pu Hoat, Nghe An Province, Vietnam (Yuan et al. 2011; Nishikawa et al. 2013 b; this study; Fig. 2).

Tylototriton notialis Stuart, Phimmachak, Sivongxay & Robichaud 2010 Tylototriton asperrimus (part) Yuan, Jiang, L��, Yang, Nguyen, Nguyen, Jin & Che 2011: 583. Tylototriton notialis Nishikawa, Matsui & Nguyen 2013 b: 36. Referred material. FMNH 271120 (male paratype), FMNH 271121 (male holotype), FMNH 271125 (one larva), FMNH 271129 (nine larvae), Laos, Khammouan Province, Boualapha District, Nakai-Nam Theun National Protected Area, Nam On River catchment, Phou Ak escarpment, 17 �� 38 ��� 39.6 ���N 105 �� 44 ��� 12.3 ���E, 980 m elev., coll. 26���29 May 2007 by Bryan L. Stuart, Somphouthone Phimmachak and Niane Sivongxay. FMNH 271122 (female paratype), same data as holotype except 17 �� 39 ���03.4���N 105 �� 44 ��� 25.2 ���E, ca. 1,000 m elev., coll. 22 May 2006 by William G. Robichaud. NUOL 0 0 443 (one topotype female), same data as types except 17.64514 ��N 105.73681 ��E, 984 m elev., coll. 14 May 2013 by Somphouthone Phimmachak, Bryan L. Stuart, and Jennifer A. Sheridan (Figs. 4 A���B). Measurements are summarized in Table 4. Distribution and natural history. This is the second report of the species in Laos, but from the same (and only) known locality in Laos. The types were collected day and night in semi-evergreen mixed with pine forest, on a stream bottom or the forest floor (Stuart et al. 2010). The larvae were found at night on the bottom of a 3 m wide stream with slow current, sand and bedrock substrate, and leaf litter-filled potholes (Stuart et al. 2010). The newly reported NUOL topotype was taken at 1115 h after heavy rain under leaf litter and pine needles at the base of a 0.3 m diameter-at-breast-height (DBH) pine tree, 4 m from a stream pool, in semi-evergreen mixed with pine forest. Elsewhere, the species has been reported from adjacent Pu Hoat, Nghe An Province, Vietnam (Yuan et al. 2011; Nishikawa et al. 2013 b; this study; Fig. 2)., Published as part of Phimmachak, Somphouthone, Aowphol, Anchalee & Stuart, Bryan L., 2015, Morphological and molecular variation in Tylototriton (Caudata: Salamandridae) in Laos, with description of a new species, pp. 285-310 in Zootaxa 4006 (2) on pages 296-297, DOI: 10.11646/zootaxa.4006.2.3, http://zenodo.org/record/237189, {"references":["Stuart, B. L., Phimmachak, S., Sivongxay, N. & Robichaud, W. G. (2010) A new species in the Tylototriton asperrimus group (Caudata: Salamandridae) from central Laos. Zootaxa, 2650, 19 - 32.","Yuan, Z. - Y., Jiang, K., Lu, S. - Q., Yang, J. - X., Nguyen, T. Q., Nguyen, T. T., Jin, J. - Q. & Che, J. (2011) A phylogeny of the Tylototriton asperrimus group (Caudata: Salamandridae) based on a mitochondrial study: suggestions for a taxonomic revision. Zoological Research, 32, 577 - 584.","Nishikawa, K., Matsui, M. & Nguyen, T. T. (2013 b) A new species of Tylototriton from northern Vietnam (Amphibia: Urodela: Salamandridae). Current Herpetology, 32, 34 - 49. http: // dx. doi. org / 10.5358 / hsj. 32.34"]}

Tylototriton panhai Nishikawa, Khonsue, Pomchote & Matsui 2013 a Tylototriton verrucosus (part) Wongratana 1984: 107. Tylototriton verrucosus Type II Pomchote, Pariyanonth & Khonsue 2008: 39. Referred material. NCSM 82961 (one female), NUOL 0 0 437 (one male), Laos, Xaignabouli Province, Botene District, Borsan Village, Phou Samrium Mountain, 17.67194 ��N 101.03081 ��E, 1,596 m elev., coll. 25���26 July 2011 by Somphouthone Phimmachak and Sengvilay Seateun. NCSM 82954 ��� 60 (seven males), same data as NCSM 82961 except 17.67656 ��N 101.03214 ��E. NUOL 0 0 421 (one female), NUOL 00422��� 28 (seven males), NUOL 0 0 452 (egg clutch), Laos, Xaignabouli Province, Botene District, Borsan Village, Houay Long Ya Stream, 17.69086 ��N 101.04661 ��E, 1,347 m elev., coll. 26 July 2011 by Somphouthone Phimmachak and Sengvilay Seateun (Fig. 5). Measurements are summarized in Table 6. Distribution and natural history. All were found in water, with some in a 2 x 4 m pond in montane shrub forest at 1,596 m elev., and some in a small sandy stream in semi-evergreen forest at 1,347 m elev. The eggs were adhered to sticks slightly above the surface of the stream (Fig. 5 D). This is the first report of the species in Laos. Elsewhere it occurs in Phu Hin Rong Kla National Park, Phitsanulok Province, Phu Luang Wildlife Sanctuary, Loei Province, and Phu Suan Sai National Park, Loei Province, Thailand (Nishikawa et al. 2013 a; Fig. 2)., Published as part of Phimmachak, Somphouthone, Aowphol, Anchalee & Stuart, Bryan L., 2015, Morphological and molecular variation in Tylototriton (Caudata: Salamandridae) in Laos, with description of a new species, pp. 285-310 in Zootaxa 4006 (2) on pages 299-300, DOI: 10.11646/zootaxa.4006.2.3, http://zenodo.org/record/237189, {"references":["Nishikawa, K., Khonsue, W., Pomchote, P. & Matsui, M. (2013 a) Two new species of Tylototriton from Thailand (Amphibia: Urodela: Salamandridae). Zootaxa, 3737 (3), 261 - 279. http: // dx. doi. org / 10.11646 / zootaxa. 3737.3.5","Wongratana, T. (1984) Range extension of the Crocodile Salamander, Tylototriton verrucosus, to Phu Luang, Thailand. Natural History Bulletin of the Siam Society, 32, 107 - 110.","Pomchote, P., Pariyanonth, P. & Khonsue, W. (2008) Two distinctive color patterns of the Himalayan Newt Tylototriton verrucosus (Urodela: Salamandridae) found in Thailand and its implication on geographic separation. The Natural History Journal of Chulalongkorn University, 8, 35 - 43."]}

Understanding introduction routes for wildlife pathogens is vital for the development of threat abatement plans. The chytrid fungus Batrachochytrium salamandrivorans (Bsal) has recently emerged in Europe, where it is considered to be a serious threat for urodelan conservation. If the highly diverse Chinese urodelans were to constitute a Bsal reservoir, then the significant international trade in these species may vector Bsal into naive urodelan communities. Here, we analyzed a total of 1,143 samples, representing 36 Chinese salamander species from 51 localities across southern China for the presence of Bsal. We found Bsal was present across a wide taxonomic, geographical, and environmental range. In particular, Bsal DNA was detected in 33 samples from the genera Cynops, Pachytriton, Paramesotriton, Tylototriton, and Andrias, including the heavily traded species Paramesotriton hongkongensis and Cynops orientalis. The true Bsal prevalence across our data set was estimated between 2% and 4%, with a maximum of 50% in a population of P. hongkongensis. Even at this overall relatively low Bsal prevalence, the exportation of millions of animals renders Bsal introduction in naive, importing countries a near certainty, which calls for the urgent implementation of proper biosecurity in the international wildlife trade.

Phylogenetic relationships of members of the salamander family Salamandridae were examined using complete mitochondrial genomes collected from 42 species representing all 20 salamandrid genera and five outgroup taxa. Weighted maximum parsimony, partitioned maximum likelihood, and partitioned Bayesian approaches all produce an identical, well-resolved phylogeny; most branches are strongly supported with greater than 90% bootstrap values and 1.0 Bayesian posterior probabilities. Our results support recent taxonomic changes in finding the traditional genera Mertensiella, Euproctus, and Triturus to be non-monophyletic species assemblages. We successfully resolved the current polytomy at the base of the salamandrid tree: the Italian newt genus Salamandrina is sister to all remaining salamandrids. Beyond Salamandrina, a clade comprising all remaining newts is separated from a clade containing the true salamanders. Among these newts, the branching orders of well-supported clades are: primitive newts (Echinotriton, Pleurodeles, and Tylototriton), New World newts (Notophthalmus-Taricha), Corsica-Sardinia newts (Euproctus), and modern European newts (Calotriton, Lissotriton, Mesotriton, Neurergus, Ommatotriton, and Triturus) plus modern Asian newts (Cynops, Pachytriton, and Paramesotriton).Two alternative sets of calibration points and two Bayesian dating methods (BEAST and MultiDivTime) were used to estimate timescales for salamandrid evolution. The estimation difference by dating methods is slight and we propose two sets of timescales based on different calibration choices. The two timescales suggest that the initial diversification of extant salamandrids took place in Europe about 97 or 69Ma. North American salamandrids were derived from their European ancestors by dispersal through North Atlantic Land Bridges in the Late Cretaceous ( approximately 69Ma) or Middle Eocene ( approximately 43Ma). Ancestors of Asian salamandrids most probably dispersed to the eastern Asia from Europe, after withdrawal of the Turgai Sea ( approximately 29Ma).

Tylototriton shanjing Nussbaum, Brodie & Yang, 1995 was described as a distinct species hidden before under the widely distributed T. verrucosus Anderson, 1871. Therefore, papers published before 1995, including those on reproductive biology, could not distinguish between these two close relatives. Consequently, data on the reproductive biology of T. shanjing are scarce. Here, we report on the first captive breeding of T. shanjing in the Aquarium of the Cologne Zoo, and we document the ontogenetic developmental stages and describe the larva for the first time. Moreover, we give a literature review for both T. verrucosus and T. shanjing and compare our data with published information referring undoubtedly to the latter species. To avoid future misidentifications and to facilitate the composition of breeding groups, we present an identification key for the East Asian salamandrid genera in general, and for the species of Tylototriton in particular.

Synopsis A fossil salamandrid, Carpathotriton matraensis gen. et sp. nov., from the Astaracian (Middle Miocene, MN 7) localities of Matraszolos 1 and 2 (Northern Hungary, Nograd County) is described. Within newts, Carpathotriton is diagnosed by a combination of primitive and derived characters including a premaxillary‐frontal contact, a well‐developed fronto‐squamosal arch, low supraorbital crest and weak sculpture on the frontal surface. Referred material comprises abundant cranial, vertebral and appendicular skeletal elements that exhibit limited morphological variation of mainly allometric nature. The dentary of Carpathotriton is unique in displaying a prominent dental parapet with an extremely short tooth row and a relatively long toothless posterior ramus. The quadrate is provided with an extuberant anterolateral process that may be a homoplasy with some members of the Tylototriton group. Trunk vertebrae of Carpathotriton have extremely high neural spines with some enlargement on their dorsal margin ...

To reconsider a classic theory that prolactin induces water-drive behavior in migratory salamanders (i.e., prolactin-based water-drive theory), I showed, with a brief review of past references, some disprovable data on the basis of “migratory activity type” and “mucus secretion” in the hynobiid salamanders, Hynobius hidamontanus and Salamandrella keyserlingii. This theory could not explain fall immigrations toward terrestrial hibernacula unrelated to mating in S. keyserlingii. Likewise, literature review indicated that it also could not explain breeding immigrations toward dry pond basins in Ambystoma opacum and toward terrestrial nest-sites adjacent to the water in Tylototriton (Echinotriton) andersoni. Mucus that was secreted from dermal mucous glands, a target organ for prolactin, was absent on the skin of both fall-nonbreeding and spring-breeding immigrants of S. keyserlingii. In the context of existence of ecological and physiological examples against the prolactin-based water-drive theory, ...

The entire mitogenome of Tylototriton kweichowensis is 16 727 bp in length. It consists of 13 protein-coding genes (PCGS), 2 ribosomal RNA genes (rRNA), 22 transfer RNA genes (tRNA), and 1 control region (CR). Except for ND6 subunit and 8 tRNA genes are distributed on the L-strand, all the other PCGs and tRNA genes are located on the H-strand. "ATG" and "GTG" are the start codons of the PCGs, "TAA", "AGA", "TA-" and "T--" are the stop codons. Most of the tRNA genes can be folded into typical clover-leaf secondary structure. The genome of T. kweichowensis has two repeat sequences in the cob-noncoding region. Mitogenomic phylogenetic analysis (NJ tree) robustly resolved the genus-level relationship among the three genera Tylototriton, Echinotriton, and Pleurodeles, and which is congruent with the previous molecular phylogeny results.

The complete mitogenome of Tylototriton shanjing is 16,661 bp in length with GenBank accession number KR154461, which contains 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNA), 2 ribosomal RNA genes (rRNA), and 1 control region (CR). The overall base composition of this mitogenome is biased toward AT content at 59.45%. Most of the PCGs and tRNA genes are located on the H-strand, except for ND6 subunit gene and eight tRNA genes, which were distributed on the L-strand. The PCGs used “ATG” and “GTG” as the start codons, while “TAA”, “TAG”, “AGA”, and “T–” are used as stop codons. Almost all tRNA genes were folded into typical cloverleaf secondary structures. The T. shanjing genome had two tandem repeat sequences in the cob-noncoding region. The mitogenomic phylogenetic analyses shows that the genera Echinotriton and Tylototriton were clustered into a strong supported monophyletic clade, which is a sister clade to the genus Pleurodeles, this confirms the previous phylogenetic results.

We examine phylogenetic relationships among salamanders of the family Salamandridae using approximately 2700 bases of new mtDNA sequence data (the tRNA(Leu), ND1, tRNA(Gln), tRNA(Gln), tRNA(Met), ND2, tRNA(Trp), tRNA(Ala), tRNA(Asn), tRNA(Cys), tRNA(Tyr), and Cot genes and the origin for light-strand replication) collected from 96 individuals representing 61 of the 66 recognized salamandrid species and outgroups. Phylogenetic analyses using maximum parsimony and Bayesian analysis are performed on the new data alone and combined with previously reported sequences from other parts of the mitochondrial genome. The basal phylogenetic split is a polytomy of lineages ancestral to (1) the Italian newt Salamandrina terdigitata, (2) a strongly supported clade comprising the "true" salamanders (genera Chioglossa, Mertensiella, Lyciasalamandra, and Salamandra), and (3) a strongly supported clade comprising all newts except S. terdigitata. Strongly supported clades within the true salamanders include monophyly of each genus and grouping Chioglossa and Mertensiella as the sister taxon to a clade comprising Lyciasalamandra and Salamandra. Among newts, genera Echinotriton, Pleurodeles, and Tylototriton form a strongly supported clade whose sister taxon comprises the genera Calotriton, Cynops, Euproctus, Neurergus, Notophthalmus, Pachytriton, Paramesotriton, Taricha, and Triturus. Our results strongly support monophyly of all polytypic newt genera except Paramesotriton and Triturus, which appear paraphyletic, and Calotriton, for which only one of the two species is sampled. Other well-supported clades within newts include (1) Asian genera Cynops, Pachytriton, and Paramesotriton, (2) North American genera Notophthalmus and Taricha, (3) the Triturus vulgaris species group, and (4) the Triturus cristatus species group; some additional groupings appear strong in Bayesian but not parsimony analyses. Rates of lineage accumulation through time are evaluated using this nearly comprehensive sampling of salamandrid species-level lineages. Rate of lineage accumulation appears constant throughout salamandrid evolutionary history with no obvious fluctuations associated with origins of morphological or ecological novelties. (c) 2006 Elsevier Inc. All rights reserved.

MtDNA sequences (396 bp cytochrome b and 369 bp 12S rRNA) from recent material and old museum specimens indicate Pleurodeles poireti and P. waltl form independent clades with 7.76% genetic divergence. Within P. poireti, populations from Djebel Edough, NE Algeria are very distinct with 6.12% genetic divergence from the remainder and may deserve separate species status. Away from Djebel Edough, P. poireti consists of three distinct clades (coastal NW Tunisia; central N Algeria; Constantine plus inland NW Tunisia) with a maximum genetic divergence of only 1%. P. waltl contains two clades with 2.96% genetic divergence, one in SE and E Spain plus north Morocco, the other in Portugal and SW and central Spain. Pleurodeles probably invaded NW Africa from SW Europe during the Messinian Salinity Crisis, when land contact was first established at 5.6 Ma, and then interrupted at 5.3 Ma. Molecular clocks, calibrated in the assumption that the latter event separated P. waltl and P. poireti, suggest that Pleuro...

In this paper, the complete 16,725 bp nucleotide sequence of the mitochondrial genome was determined for the Tylototriton kweichowensis (Caudata:Salamandridae). It contains 37 genes (13 protein-coding genes, 2 rRNA genes and 22 tRNA genes) and a non-coding region (D-loop). Overall base composition of the complete mitochondrial DNA is A (33.8%), G (14.4%), C (26.2%), and T (25.6%), so the percentage of A and T (59.4%) is higher than G and C (40.6%). All the genes in T. kweichowensis are distributed on the H-strand, except for the ND6 subunit gene and eight tRNA genes which are encoded on the L-strand.

We observed male courtship behavior of Tylototriton (Echinotriton) andersoni in the laboratory. In some males, the cloaca swelled and became wet with mucous secretions from December to May. A male in this condition crept around a female, sniffing around her body and drawing a thread of mucus from his cloaca, so that the female was surrounded by spiderweb-like string of mucous attached to the substrate. The male then deposited a spermatophore by rubbing his cloaca against the substratum while swaying his body to and fro. The male behaved similarly towards females from different localities, though we could not observe any female reaction to the male. Under a phase-contrast microscope, the gross morphology of sperms from a male from Amamioshima Island was similar to that reported for males from Okinawajima.

The Chinhai Salamander, Echinotriton chinhaiensis, mates on land. The male of a courting pair marks a trail by a mucous excretion from his cloaca. The pair follows an ellipsoid track during which manoeuvre a spermatophore is deposited. Partners align their bodies such that the female is led over the spermatophore. Clasping was not observed. The female traces the spermatophore with her cloaca through shifting movements with the pelvic region. A male may deposit several spermatophores on one evening. The courtship is compared to that in the genus Tylototriton.

Tylototriton uyenoi sp. nov. (Figs. 1 B, 6 A, B, G, and J) Tylototriton verrucosus: Smith (1924): 309; Taylor (1962): 279; Tylototriton verrucosus Type I: Pomchote et al. (2008): 39 (part). Identity: This species corresponds to the Group 2 (T. shanjing Lineage 4 in molecular analyses). Holotype: KUHE 19147, an adult male from Phuping Rajanives Palace, Doi Suthep, Chiang Mai Province, Thailand (18 r 48 ��� 16 ������ N, 98 r 54 ��� 9 ������ E, 1436 m asl), collected on 10 August 1994 by Masafumi Matsui and Jarujin Nabhitabhata. Paratypes: A total of eight males: KUHE 19037 and 19038 from Royal Garden Siribhume, Doi Inthanon, Chiang Mai Province, Thailand (18 r 32 ��� 46 ������ N, 98 r 31 ��� 14 ������ E, 1313 m asl), collected on 1 August 1994 by Masafumi Matsui and Jarujin Nabhitabhata, KUHE 19146, and 19148 ��� 50 (data same as the holotype), and NSMT- H 1076 and 1077 from Doi Suthep, Chiang Mai Province, Thailand, collected on 8 September 1967 by Shun-ichi U��no; two females: NSMT-H 1073 and 1075 (data same as NSMT-H 1076 and 1077); a larva: NSMT-H 1074 (data same as NSMT-H 1076 and 1077). Etymology: The specific epithet is dedicated to Dr. Shun-ichi U��no, who collected part of the paratypes of the new species. Diagnosis: The new species is placed in the genus Tylototriton by having the combination of: dorsal warts present; dorsolateral bony ridges on head present; knob-like warts on dorsolateral body present; quadrate spine absent. A medium-sized newt of Tylototriton; skin rough with fine granules; dorsolateral bony ridges on head prominent and narrow; vertebral ridge distinct and slightly segmented; rib nodules prominent; limbs long and thin; tips of forelimbs and hindlimbs greatly overlapping when adpressed along body; tail thin; dorsal head, upper and lower lips, vertebral ridge, rib nodules, limbs, vent region, and whole tail orange to reddish brown. Description of holotype: Body moderately stout; skin rough; fine granules dense on dorsum, arranged in transverse striations on mid ventrum, and small and sparse on throat; head wider than long, hexagonal in shape, depressed and slightly oblique in profile; snout short, truncate, slightly beyond lower jaw; nostril close to snout tip; dorsolateral bony ridges on head prominent, narrow, sharply protruding, and slightly rough, from above eye to above anterior end of parotoid, posterior ends scrolled proximally; middorsal ridge on head short, weak; labial fold absent; skull wide and hexagonal, dorsolateral crests steep and narrow; middorsal crest prominent; maxillary connecting with quadrate, but slightly separated from pterygoid; front-squamosal arch robust and wide; vomerine tooth series in an inverted V-shape, converging anteriorly and reaching choana; tongue oval and attached to mouth floor but free laterally and posteriorly; parotoid distinct but small, and projecting posteriorly; gular fold present; costal folds absent; vertebral ridge prominent and slightly segmented, from neck to base of tail, separated from middorsal ridge on head with a small gap; rib nodules distinct but small, forming knob-like warts, 15 on right side and 14 on left side of body from axilla to base of tail; rib nodules slightly increasing in size from most anterior to fourth nodule, then decreasing posteriorly; tips of fore- and hindlimbs greatly overlapping when adpressed along body (longest toe reaching at elbow); number of trunk vertebrae (presacral vertebrae including atlas) 14; fingers and toes free of webbing; tail laterally compressed, dorsal fin more distinct posteriorly, ventral edge smooth, tip pointed; numbers of upper jaw teeth 107, lower jaw teeth 119, and vomerine teeth 102. Measurements of holotype (in mm): SVL 68.6; HL 15.9; HW 16.5; MXHW 17.7; SL 6.4; LJL 14.0; ENL 3.9; IND 4.8; IOD 7.4; UEW 2.3; UEL 5.0; OL 3.3; AGD 34.8; TRL 52.7; TAL 87.6; VL 6.2; BTAW 5.5; MTAW 3.4; BTAH 8.5; MXTAH 9.0; MTAH 7.5; FLL 24.8; HLL 28.9; 2 FL 4.6; 3 FL 5.0; 3 TL 6.8; 5 TL 3.3; VTW 5.1; VTL 6.9. Color: In life, dorsal ground color dark brown to black; venter slightly lighter than dorsum; anterior half of head, parotoids, vertebral ridge, rib nodules, limbs, vent region, and whole tail orange to reddish brown. In preservative, dorsal coloration tending to fade, becoming light brown and orange to brown markings fading to cream. Variation: Males have more robust limbs than females. Males have a relatively longer tail (RTAL: 101.4��� 127.7, median=115.0 %SVL in males vs. 88.0 and 97.0%SVL in females) and vent slit (RVL: 6.0���9.0, median= 7.4 %SVL in males vs. 1.7 and 1.9 %SVL in females) than females. Male specimens of the type series are basically similar in morphology, but some specimens have more distinctly segmented vertebral ridges, larger rib nodules, and rougher dorsolateral bony ridges on head than the holotype. Length of vomerine tooth rows is variable among specimens, but reaching/exceeding posterior end of choana in all specimens except for one (KUHE 19148). Number of trunk vertebrae is always 14. Specimens are generally uniform in color pattern, but markings slightly vary in color from orange to reddish brown. One specimen (KUHE 19148) lacked distal half of right forelimb. Egg: The mean diameter of ripe ova in ovaries of the paratype (NSMT-H 1075) ranged from 1.9 to 2.8 mm (n= 6, mean= 2.4 mm). The clutch size is unknown. The animal pole is dark brown and the remaining area is dark cream in color. Larva: A paratype larva (NSMT-H 1074) was collected in Doi Suthep in early September. It is fully-grown and has a total length of 66.1 mm. The gills started to shrink. The following description of larva is based on this preserved individual. Head rounded triangle; depressed and sloping in profile; snout short and rounded; labial fold distinct at posterior half of upper jaw; caudal and dorsal fins nearly absorbed; tail tip rounded. Dorsum yellowish brown; venter paler; anterior head, parotoids, vertebral ridge, rib nodules, limbs, and tail yellow. Smith (1924) reported that the larvae at early developmental stages possessed a pair of balancers. Comparisons: Tylototriton uyenoi ( the Group 2 [T. shanjing Lineage 4]) is a member of the subgenus Tylototriton based on the molecular phylogeny. This species differs from members of the subgenus Yaotriton except for T. panhai ( the Group 3 [T. shanjing Lineage 6]), another new species described below, and T. daweishanensis, T. taliangensis, and T. v. verrucosus [data of T. daweishanensis are taken from Zhao et al. 2012 and those of T. v. verrucosus are taken from Anderson (1871)] of the subgenus Tylototriton, by having distinct orange to reddish brown markings on head, trunk, limbs, and tail (vs. black body except for ventral edge of tail in the remaining species of the subgenus Yaotriton except for T. panhai, T. daweishanensis, and T. v. verrucosus, and black body except for posterior parotoids in T. taliangensis). The new species differs from T. kweichowensis and T. pseudoverrucosus by having isolated orange to reddish brown markings on rib nodules (vs. connected markings forming dorsolateral lines in T. kweichowensis and T. pseudoverrucosus [data of T. pseudoverrucosus are taken from Hou et al. 2012]), from T. shanjing by having darker markings, relatively wider head, longer and higher tail, and wider and longer vomerine teeth series (vs. bright markings, narrower head, shorter and lower tail, and narrower and shorter vomerine teeth series in T. shanjing), from T. panhai by having orange to reddish brown limbs and tail, and narrow dorsolateral ridges on head (vs. black limbs and tail except for edges, and wide dorsolateral ridges on head in T. panhai), from T. v. pulcherrima by having black ground color of the body (vs. light brown ground color in T. v. pulcherrima), from T. yangi by having orange anterior half of head (vs. black in T. yangi). Range: Doi Ang Khang, Doi Chang Kien, Doi Inthanon, Doi Pui, and Doi Suthep, Chiang Mai Province, Thailand. Natural history: The holotype and the paratypes deposited in KUHE were collected in artificial pools. The species is also found in natural or artificial ponds, ditches along farms, and in slow streams, with an average depth of 38.2 cm (range 9���120 cm). Breeding occurs from May to July. Larvae are found in the water bodies from August to December. The adults are probably semiaquatic, which are found in water even out of breeding season., Published as part of Nishikawa, Kanto, Khonsue, Wichase, Pomchote, Porrawee & Matsui, Masafumi, 2013, Two new species of Tylototriton from Thailand (Amphibia: Urodela: Salamandridae), pp. 261-279 in Zootaxa 3737 (3) on pages 272-275, DOI: 10.11646/zootaxa.3737.3.5, http://zenodo.org/record/285235

Tylototriton panhai sp. nov. (Figs. 1 C, 6 C, D, H, and K) Tylototriton verrucosus: Wongratana (1984): 107; Tylototriton verrucosus Type II: Pomchote et al. (2008): 39. Identity: This species corresponds to the Group 3 (T. shanjing Lineage 6 in molecular analyses). Holotype: THNHM 2800, an adult female from Phu Ruea, Phu Luang Wildlife Sanctuary, Loei Province, Thailand (17 r 29 ��� 59 ������ N, 101 r 20 ��� 30 ������ E, 1183 m asl), collected on 25 May 1998 by Tanya Chan-ard. Etymology: The specific epithet is dedicated to Prof. Somsak Panha (Chulalongkorn University), who is an active naturalist and helped our field surveys. Diagnosis: The new species is placed in the genus Tylototriton by having the combination of: dorsal warts present; dorsolateral bony ridges on head present; knob-like warts on dorsolateral body present,; quadrate spine absent. A medium-sized Tylototriton; skin rough with fine granules; dorsolateral bony ridges on head wide; vertebral ridge distinct and not segmented; rib nodules prominent and large; limbs long and thin; tips of forelimbs and hindlimbs greatly overlapping when adpressed along body; tail thin; dorsal head, upper and lower lips, parotoids, vertebral ridge, rib nodules, tips of fingers and toes, margin of vent slit, and dorsal and ventral edges of tail yellow, orange, to reddish brown. Description of holotype: Body moderately stout; skin rough; fine granules dense on dorsum, arranged in transverse striations on mid ventrum, and small and sparse on throat; head longer than wide, hexagonal in shape, depressed and slightly oblique in profile; snout short, truncate, slightly beyond lower jaw; nostril close to snout tip; dorsolateral bony ridges on head prominent, wide, moderately protruding, and slightly rough, from above eye to above anterior end of parotoid, posterior ends slightly scrolled proximally; middorsal ridge on head absent; labial fold absent; skull wide and hexagonal, dorsolateral crests moderate and wide, middorsal crest absent; maxillary connecting with quadrate, but slightly separated from pterygoid; front-squamosal arch robust and wide; vomerine tooth series in an inverted V-shape, converging anteriorly and not reaching posterior end of choana; tongue oval and attached to mouth floor but free laterally and posteriorly; parotoid large, protruding dorsolaterally and posteriorly; gular fold present; costal folds absent; vertebral ridge prominent and smooth, from neck to base of tail; rib nodules distinct and large except for the anteriormost one on right side, forming knob-like warts, 14 on both sides of body from axilla to base of tail; rib nodules slightly increasing in size from most anterior to fourth or fifth nodule, then decreasing posteriorly; tips of fore- and hindlimbs overlapping when adpressed along body (longest toe reaching distal part of palm); number of trunk vertebrae 14; fingers and toes free of webbing; tail laterally compressed, dorsal fin more distinct posteriorly, ventral edge smooth, tip pointed; numbers of upper jaw teeth 80, lower jaw teeth 81, and vomerine teeth 66. Measurements of holotype (in mm): SVL 71.6; HL 17.6; HW 16.9; MXHW 18.9; SL 7.4; LJL 13.8; ENL 3.9; IND 5.4; IOD 7.6; UEW 2.5; UEL 4.1; OL 2.9; AGD 36.5; TRL 54.0; TAL 63.3; VL 3.0; BTAW 5.9; MTAW 3.9; BTAH 8.9; MXTAH 9.0; MTAH 7.6; FLL 20.2; HLL 22.4; 2 FL 3.2; 3 FL 3.5; 3 TL 6.0; 5 TL 2.7; VTW 4.0; VTL 6.8. Color: In life, dorsum ground color dark brown; venter slightly lighter than dorsum; anterior half of head, parotoids, vertebral ridge, rib nodules, tips of fingers and toes, margin of vent slit, and dorsal and ventral edges of tail yellow, orange, to reddish brown (based on the unvouchered samples whose tail tips were used for DNA analysis). In preservative, dorsal coloration tending to fade, becoming light brown and orange to brown markings fading to cream (the holotype). Variation: Our field observation found a variation in color of markings, from yellow, dark orange, to reddish brown, and in size of rib nodules. Individuals from Phu Hin Rong Kla National Park, Phitsanulok Province have dark reddish brown markings, and those from Phu Suan Sai National Park, Loei Province have smaller nodules than the holotype. Egg: The mean diameter of ripe ova in right ovary that could be measured by a minor dissection of the holotype ranged from 1.5 to 2.7 mm (n= 7, gross mean= 2.1 mm). The clutch size is unknown. The animal pole is dark brown and the remaining area is dark cream in color. Comparisons: Tylototriton panhai ( the Group 3 [T. shanjing Lineage 6]) is a member of the subgenus Yaotriton based on the molecular phylogeny, and differs from all other species of the subgenus by having distinct yellow to reddish brown markings on head, trunk, and tail edges (vs. having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail in all other species of Yaotriton). The species looks similar to most of the species of the subgenus Tylototriton, especially T. pseudoverrucosus, T. shanjing, T. uyenoi ( the Group 2 [T. shanjing Lineage 4]), and T. v. pulcherrima, however differs by having widely developed dorsolateral bony ridges on head, black limbs and tail except for ridges (vs. less developed dorsolateral bony ridges on head, yellow or orange limbs and whole tail in T. pseudoverrucosus [data of T. pseudoverrucosus are taken from Hou et al. 2012], T. shanjing, T. uyenoi, and T. v. pulcherrima). The species differs from the remaining species of the subgenus Tylototriton in the following way: from T. daweishanensis by having distinct yellow to reddish brown markings on head, trunk, and tail edges (vs. having black body except for tips of fingers and toes, margin of vent, and ventral edge of tail in T. daweishanensis [data are taken from Zhao et al. 2012]), from T. kweichowensis and T. taliangensis by having isolated rib nodules (vs. connected nodules forming dorsolateral rides on body in T. kweichowensis and T. taliangensis), from T. kweichowensis and T. yangi by having colored anterior half of head (vs. black head except for posterior parotoids and jaw angles in T. kweichowensis and T. yangi), and from T. v. verrucosus by having colored markings (vs. black body in T. v. verrucosus [data of T. v. verrucosus are taken from Anderson 1871]). Range: Phu Hin Rong Kla National Park, Phitsanulok Province, and Phu Luang Wildlife Sanctuary and Phu Suan Sai National Park, Loei Province, Thailand. The locality in Phu Hin Rong Kla National Park is the southernmost record in the genus Tylototriton. Natural history: The species was first reported by Wongratana (1984), which was collected walking on an elephant trail at foggy morning. The habitat is moist forest with bamboo bushes and variety of water bodies like temporal pools by rain, and streams. Eggs are found as attached on wet grass or plant overhanging the water surface or under wood debris on land near small streams or ponds. In the breeding season from May to July, the adults were found in the water bodies. Out of the breeding season, the species is probably terrestrial and difficult to encounter., Published as part of Nishikawa, Kanto, Khonsue, Wichase, Pomchote, Porrawee & Matsui, Masafumi, 2013, Two new species of Tylototriton from Thailand (Amphibia: Urodela: Salamandridae), pp. 261-279 in Zootaxa 3737 (3) on pages 275-276, DOI: 10.11646/zootaxa.3737.3.5, http://zenodo.org/record/285235

We newly sequenced the mitochondrial genome of Tylototriton wenxianensis. The total length of the T. wenxianensis mitogenome is 16 265 bp, with GenBank accession number KR733683. It consists of 13 protein-coding genes (PCGs), two ribosomal RNA genes (rRNA), 22 transfer RNA genes (tRNA), and one control region (CR). Most of the genes are encoded on the H-strand, except for eight tRNA and ND6, which are encoded on the L-strand. Our mitogenomic phylogenetic tree showed that the relationships among the genera Tylototriton, Echinotriton, and Pleurodeles were well supported, and which is consistent with the previous molecular phylogeny.

Tylototriton taliangensis was listed as a Near Threatened amphibian in IUCN red list. In this study, we sequenced the complete mitochondrial (mt) genome of this species (GenBank: KP979646) and found it contains 16,265 base pairs, which encode 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNA), 2 ribosomal RNA genes (rRNA) and 1 control region (CR). We also found that almost all PCGs and tRNA genes are located on the H-strand, except for ND6 subunit gene and eight tRNA genes, which were distributed on the L-strand. The PCGs used "ATG" and "GTG" as the start codon, while used four types of stop codons. Almost all tRNA genes were folded into typical cloverleaf secondary structures. The L-strand replication origin (OL) and a non-coding region were also found. The new mitogenomic phylogenetic tree confirms the reciprocally monophyly of the genus Tylototriton, Echinotriton and Pleurodeles with high bootstrap value. The present study will provide information for future studies on the conservation genetics and phylogeny of this species and its relatives.

Until now, Asian amphibians appear to have largely escaped declines driven by chytridiomycosis. Vietnamese salamanders that belong to the genus Tylototriton are rare and have a patchy distribution in mountainous areas, falling within the proposed environmental envelope of chytrid infections, surrounded by Batrachochytrium dendrobatidis infected regions. If these salamanders are susceptible to chytridiomycosis, then their populations could be highly vulnerable after the introduction of B. dendrobatidis. Examination for the presence of the chytrid fungus in skin swabs from 19 Tylototriton asperrimus and 104 Tylototriton vietnamensis by using quantitative polymerase chain reaction was performed. Susceptibility of T. asperrimus to experimental infection by using the global panzootic lineage (BdGPL) strain of B. dendrobatidis was examined. The fungus was absent in all samples from all wild salamanders examined. Inoculation with the BdGPL strain resulted in mortality of all five inoculated salamanders within 3 weeks after inoculation with infected animals that manifested severe orthokeratotic hyperkeratosis, epidermal hyperplasia, and spongiosis. Although infection by B. dendrobatidis currently appears absent in Vietnamese Tylototriton populations, the rarity of these animals, their pronounced susceptibility to chytridiomycosis, an apparently suitable environmental context and increasing likelihood of the pathogen being introduced, together suggest the need of urgent measures to avoid future scenarios of extinction as witnessed in Central America and Australia.

The skull of neotenic individuals of the Alpine Newt Triturus alpestris from the locality Drakolimni (Greece) is described on the basis of models made from magnified serial frontal sections. In order to recognize features associated with neoteny and paedomorphosis, the results were compared with normal development of contemporary Triturus and of other Caudata. The neotenic larvae from Greece correspond to advanced stages of metamorphosis in normal development of Triturus alpestris. Comparison with salamandrids from the Tertiary of Europe in which the hyobranchial skeleton was preserved, namely Brachycormus noachicus, Chelotriton paradoxus, and Palaeopleurodeles hauffi, revealed that both latter taxa were completely metamorphosed adults anatomically similar to their contemporary relatives Tylototriton, Echinotriton, and Pleurodeles, whereas Brachycormus, though apparently related to Chelotriton, was a neotenic amphibian. This is suggested by its incompletely metamorphosed but ossified hyobranchial skeleton. © 1996 Wiley-Liss, Inc.

Tylototriton notialis sp. nov. Holotype: FMNH 271121 (field tag BLS 10775), adult male (Fig. 3), Laos, Khammouan Province, Boualapha District, Nakai-Nam Theun National Protected Area, Nam On River catchment, Phou Ak escarpment, 17 �� 38 ' 39.6 "N 105 �� 44 ' 12.3 "E (Fig. 4), 980 m elev., coll. 27 May 2007 by Bryan L. Stuart, Somphouthone Phimmachak, and Niane Sivongxay. Paratypes: FMNH 271120, adult male (Fig. 3), same data as holotype except coll. 26 May 2007. FMNH 271122, adult female, same data as holotype except 17 �� 39 '03.4"N 105 �� 44 ' 25.2 "E, ca. 1,000 m elev., coll. 22 May 2006 by William G. Robichaud. Referred material: FMNH 271125 (one larva; Fig. 5), same data as paratype male. FMNH 271129 (nine larvae), same data as paratype male except coll. 29 May 2007. Subgenus: The new species is assigned to the subgenus Yaotriton Dubois & Raffa��lli 2009 (= T. asperrimus species group of Fei et al. 2005) based on its phylogenetic position (Fig. 1) and mostly dark dorsal and ventral coloration (Dubois & Raffa��lli 2009). Etymology: The specific epithet taken from notialis L. for southern, in reference to the new species having the southernmost known locality of any member in the T. asperrimus group. Diagnosis: Tylototriton notialis is a Tylototriton (Yaotriton) having the combination of distinct, knob-like rib nodules (dorsolateral row of glandular warts); glandular warts on most of the remaining dorsal and ventral surfaces; very dark brown to black coloration on dorsum and venter; and bright orange coloration on rib nodules and posterior end of parotoid. Description of holotype: Habitus moderately stout. Head broader than body, longer than wide, slightly sloping in profile. Snout short, truncate in dorsal view, rounded in profile, exceeding beyond lower jaw. Nostrils close to snout tip, right nostril slightly visible from above. Vomeropalatine teeth in two rows, anteriorly in contact and beginning posterior to the anterior margin of the choanae, converging into two parallel rows briefly, then diverging from one another. Glandular ridge on midline of crown from above anterior edge of eye to middle of head. Glandular ridge on outer margin of crown from above eye to base of parotoid. Parotoids enlarged, projecting backward. Glandular patch of skin on nape. Distinct vertebral tubercular ridge from posterior end of crown to base of tail, separated from ridge on midline of crown. Dorsolateral row of approximately 14 large glandular warts (rib nodules) on each side from level of axilla to base of tail, distinctly knob-like anteriorly, becoming smaller posteriorly, merging at level of groin. Smaller, glandular warts on most of remaining dorsal and ventral surfaces, warts on crown, nape, and back with clusters of glands and sometimes conical, those on throat granular and widely spaced, those on belly arranged in striations perpendicular to body axis, with a smooth, glandular, ovoid patch of skin on chest. Weak gular fold present. Four fingers, five toes, all without webbing. Tail laterally compressed, narrow dorsal fin, smooth ventral ridge, tip acuminate in profile. Color of holotype in life: Body very dark brown. Margin of upper and lower lip, posterior end of parotoid, dorsolateral glands (rib nodules), dorsal margin of tail, outer margin of upper surface of hand and finger tips, lower surface of hand except center of palm, dorsal surface of toe tips, ventral surface of toes and extending from fifth toe to base of foot, cloacal region continuing to ventral ridge of tail bright orange. Anterior surface of forelimb from axilla to base of first finger with narrow, broken, yellow stripe. Anterior and ventral surfaces of hindlimbs with small, scattered yellow spots. Iris dark brown, pupil black. Color of holotype in preservative: Body faded to a lighter shade of brown. Orange markings faded to yellow or creamy-yellow. Measurements of holotype: SVLA 60.2; SVLP 69.1; AXGR 30.5; TTL 130.4; TAL 63.1; TAD 6.5; HL 19.6; HW 15.9; EN 4.0; IN 4.9; AL 23.7; PL 23.0. Variation: The body coloration of the paratype male (collected on land) was black rather than very dark brown as in the holotype (collected in water) and paratype female (collected on land). B��hme et al. (2005) reported that T. vietnamensis taken on land were darker in coloration than those living in water. Measurements are summarized in Table 3. Larvae: Two size classes of larvae were obtained in syntopy with the holotype: FMNH 271125 (one larva) with TTL 33.1, and FMNH 271129 (nine larvae) with TTL 17.8���20.9 (mean 19.4 �� S.D. 0.9). Distribution and natural history: The new species is currently known only from the type locality, although it likely also occurs in adjacent Vietnam given its proximity to the border. All were taken in evergreen mixed with deciduous and pine forest (Fig. 6). The holotype and larvae were collected at 1940��� 2042 h on the stream bottom under 10���30 cm of water (pH 5.0, temperature 19.5 ��C) in a 3 m wide stream with slow current and sand and bedrock substrate, including leaf litter-filled potholes, just upstream from a 3 m high chute over bedrock. The paratype male was found at 1030 h, 30 cm above the ground on a 20 cm diameter log covered by another log, approximately 20 m from a steep escarpment. The type locality lies near the former Ho Chi Minh Trail, and a large, unexploded bomb dropped by American forces during the war was embedded in the ground approximately 3 m from the paratype male (Fig. 7). The paratype female was found during the day on the damp forest floor. The testes of the holotype and paratype males are mature but not enlarged, suggesting these males were in their first reproductive season. The paratype female has enlarged oviducts, indicating that she had recently laid eggs. Eggs of the new species were not found. Conservation: Overharvesting for traditional medicine and the international pet trade has been identified as a major threat to Asian salamandrids (Rowley et al. 2010). The formal description of Laotriton laoensis (Stuart & Papenfuss 2002), the first salamandrid known from Laos, inadvertently led to its exploitation for the international pet trade (Stuart et al. 2006). That species is now protected from commercial collecting by national legislation in Laos. It is hoped that the muted coloration and superficial similarity of T. notialis to other members of the T. asperrimus group will minimize its demand in the international pet trade. The new species��� occurrence in a remote area within Nakai-Nam Theun National Protected Area, which is one of Laos��� largest and best-funded national protected areas (WMPA 2005), should also afford it protection. Comparisons: Tylototriton notialis differs from all other species in the T. asperrimus group by having bright orange coloration on the posterior end of the parotoids (parotoid coloration dark like dorsum in all other species). Tylototriton notialis differs from T. asperrimus, T. wenxianensis, T. hainanensis, and T. cf. vietnamensis from Cao Bang Province, Vietnam, by having bright orange coloration on the rib nodules (uniformly dark dorsum in T. asperrimus, T. wenxianensis, T. hainanensis, and T. cf. vietnamensis). Tylototriton notialis differs from T. vietnamensis by having glandular warts on most of the dorsal and ventral surfaces and very dark brown to black body coloration (T. vietnamensis with mostly smooth skin and gray to tan body coloration). Tylototriton notialis further differs from T. hainanensis, T. vietnamensis, and T. wenxianensis by having knob-like rib nodules (slightly flattened rib nodules in T. vietnamensis and T. hainanensis and indistinct nodules in T. wenxianensis)., Published as part of Stuart, Bryan L., Phimmachak, Somphouthone, Sivongxay, Niane & Robichaud, William G., 2010, A new species in the Tylototriton asperrimus group (Caudata: Salamandridae) from central Laos, pp. 19-32 in Zootaxa 2650 on pages 23-27, DOI: 10.5281/zenodo.198798, {"references":["Dubois, A. & Raffaelli, J. (2009) A new ergotaxonomy of the family Salamandridae Goldfuss, 1820 (Amphibia, Urodela). Alytes, 26, 1 - 85.","Fei, L., Ye, C., Jiang, J. & Xie, F. (2005) An Illustrated Key to Chinese Amphibians. Sichuan Publishing House of Science and Technology, Chongqing, Sichuan, 340 pp.","Bohme, W., Schottler, T., Nguyen, T. Q. & Kohler, J. 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