Integrative taxonomy reveals three new taxa within the Tylototriton asperrimus complex (Caudata, Salamandridae) from Vietnam Author Bernardes, Marta Cologne Zoo, Riehler Str. 173, 50735 Cologne, Germany & Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany mrtbernardes@gmail.com Author Le, Minh Duc Faculty of Environmental Sciences, University of Science, Vietnam National University, Hanoi, 334 Nguyen Trai Road, Hanoi, Vietnam & Central Institute for Natural Resources and Environmental Studies, Vietnam National University, Hanoi, 19 Le Thanh Tong, Hanoi, Vietnam & Department of Herpetology, American Museum of Natural History, Central Park West at 79 th Street, New York, New York 10024, USA Author Nguyen, Truong Quang Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam & Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam https://orcid.org/0000-0002-6601-0880 Author Pham, Cuong The Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam & Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam Author Pham, Anh Van Faculty of Natural Science and Technology, Tay Bac University, Quyet Tam Ward, Son La City, Son La Province, Vietnam Author Nguyen, Tao Thien Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam & Vietnam National Museum of Nature, 18 Hoang Quoc Viet St., Hanoi, Vietnam Author Roedder, Dennis Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany Author Bonkowski, Michael Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany Author Ziegler, Thomas Cologne Zoo, Riehler Str. 173, 50735 Cologne, Germany & Terrestrial Ecology, Institute of Zoology, University of Cologne, Zuelpicher Str. 47 b, 50674 Cologne, Germany text ZooKeys 2020 935 121 164 http://dx.doi.org/10.3897/zookeys.935.37138 journal article http://dx.doi.org/10.3897/zookeys.935.37138 1313-2970-935-121 4D41FCF4A59544F095EFCF38D0FC127C 33263B29FD6E5F15BCFC61C058095F9C Tylototriton sparreboomi M. Bernardes, M. D. Le, T. Q. Nguyen, C. T. Pham, A. V. Pham, T.T. Nguyen & T. Ziegler sp. nov. Figures 9 , 11 Tylototriton taxon 3 (this study). T. sp.: Laking et al. 2017 , page 2. T. verrucosus : Orlov et al. 2002 , page 101; van Dijk et al. 2009 , page 1; Nguyen et al. 2009 , page 329. T. cf. asperrimus "North Vietnam": Hernandez 2018 , page 80. Holotype. IEBR 4476, adult male, collected in Sa De Phin Commune, Sin Ho District, Lai Chau Province, Vietnam, at an elevation of 1670 m a.s.l., in May 2015 by A. V. Pham and M. A. Vang. Paratypes. Two adult males, same data as the holotype: IEBR 4477 and TBU 10; two adult males, collector unknown: IEBR 4478 and IEBR 4479. Etymology. The specific epithet is dedicated to late Prof. Dr. Max Sparreboom, who has made great contributions to the understanding of Urodela . Diagnosis. The new species is distinguished from other species of the genus by the following combination of characters: head longer than wide; snout truncate in dorsal view; tips of fingers reaching nostril when foreleg is laid forward; skin tubercles on ventral side shaped like transverse wrinkles; rib nodules distinct and round; vertebral ridge segmented, high and relatively wide; relatively wide distance between the eyes; and gular and labial folds present. Description of holotype. Habitus stout; head broader than body, longer than wide, depressed and slightly oblique in profile; snout wider than long (IN> EN), truncate in dorsal view, rounded in profile and protruding beyond lower jaw; nostrils close to snout tip and not visible from above; labial fold slightly evident; dorsolateral bony ridges on head prominent, wide, moderately protruding, from above eye to above anterior end of parotoid, posterior ends slightly scrolled inside; middorsal ridge on head almost indistinct; parotoids enlarged, projecting backwards; ventral skin smoother than dorsal skin, with tubercles shaped like transverse wrinkles; gular fold weak; glandular vertebral ridge high, wide, smooth and segmented extending from top of head to base of tail, separated from middorsal ridge; number of trunk vertebrae 13; rib nodules distinct and roundish, the third anterior rib nodule on right side is located below the second nodule and the fourth nodule seems to not be associated with the fourth vertebra, nodules appear knob-like anteriorly, becoming smaller posteriorly; tips of fore- and hind limbs overlap when adpressed along body; tips of fingers reaching nostril when foreleg laid forward; and tail laterally compressed, thin and tip acuminated. Color of holotype. In preservative, with an overall faded dark brown coloration, with faded yellow markings on vent, ventral margin of tail, tips of fingers and toes, and part of palms. For color in life see Fig. 9 . Measurements of holotype (in mm). SVL 68.71; MHW 17.60; HW 9.85; HL 19.95; PL 10.18; PH 6.12; EL 3.43; EN 4.43; IN 6.26; IE 9.04; LJL 10.74; UEL 4.74; HUM 9.27; RAD 15.86; FEM 8.77; TIB 16.03; TL 59.70; TH 8.57; ClL 9.66; WVr 2.37; L5W 3.04; AG 30.99; and TkL 42.72. Variation. TBU 10 (in worse preserved condition) presents rib-nodules thinner than holotype, glandular vertebral ridge more tubercular, and tail tip slightly rounded. The remaining characters were similar to the holotype in morphology. For detailed measurements see Table 4 . Comparisons. Tylototriton sparreboomi sp. nov. differs from other related species of Tylototriton as follows: from T. anhuiensis by distinctly separated rib nodules (versus continuous nodule-like warts in T. anhuiensis ); from T. asperrimus by a head longer than wide (versus wider than long in T. asperrimus according to Nishikawa et al. 2013b ; Sparreboom 2014 ; Hernandez 2016 ), however the female holotype shows similar head proportions-see Discussion); from T. broadoridgus by head slightly longer than wide (versus equally long and wide), presence of skin on ventral side shaped like transverse wrinkles (versus covered with round shaped tubercles, like the dorsal side), distinctly separated rib nodules (versus continuous nodule-like warts), and narrower vertebral ridge (versus wider vertebral ridge in T. broadoridgus ); from T. hainanensis by a head longer than wide (versus much wider than long), tips of fingers reaching nostril (versus eyes) when foreleg is laid forward, and a snout truncate in dorsal view (versus rounded in T. hainanensis ); from T. liuyangensis by a wider (versus shorter) distance between eyes, distinctly separated rib nodules (versus continuous nodule-like warts), and lateral skin shaped like transverse wrinkles (versus covered by warts in T. liuyangensis ); from T. notialis by a broader (versus narrower) head, longer (versus shorter) hind-limbs, and higher tail (versus thinner tail in T. notialis ); from T. panhai by wider (versus shorter) distance between the eyes, presence (versus absence) of labial fold, and overall dorsal coloration mostly dark (versus with presence of characteristic dorsal colorful markings in T. panhai ); from T. pasmansi sensu lato by a narrower (versus wider) head, slightly wider (versus slightly narrower) distance between the eyes, tips of fingers reaching nostril (versus eye) when foreleg laid forward, longer (versus shorter) humerus length, and slightly enlarged round-like rib nodules (versus slightly smaller, pointy to rounded rib nodules in T. pasmansi sensu lato); from T. p. pasmansi by a longer (versus shorter) length between eye and nostril and wider (versus narrower) vertebral ridge in T. p. pasmansi ; from T. pasmansi obsti by a longer (versus shorter) femur length; from T. vietnamensis by a moderately stout (versus slender) habitus, presence (versus absence) of gular fold, and round (versus slightly flattened) rib nodules and high vertebral ridge (versus low vertebral ridge in T. vietnamensis ); from T. wenxianensis by a truncate (versus more rounded) snout in dorsal view, wider (versus narrower) distance between the eyes, distinctly separated rib nodules (versus continuous nodule-like warts), smoother (versus extremely rough) skin on ventral side shaped like transverse wrinkles (versus rounded shaped and uniform to dorsal side), and colored marking on ventral slit (versus black colored ventral slit in T. wenxianensis ); from T. ziegleri by a head longer than wide (versus wider than long), rounded but smaller (versus enlarged knob-like) rib nodules, and distinctly segmented vertebral ridge (versus even more segmented vertebral ridge in T. ziegleri ). Figure 11. Tylototriton sparreboomi sp. nov. (holotype). In sequence: dorsal view; ventral view; lateral view; and detail of dorsal view of the head. Photographs T. Ziegler. Distribution. Known only from the type locality in Lai Chau Province, northern Vietnam (Fig. 2 ). Natural history. Specimens were found in water between 9:00 and 16:30 h in ponds. The surrounding habitat was secondary forest of large, medium and small hardwoods mixed with shrubs and vines. Air temperature at the sites was 23 to 27 °C and relative humidity was 80 to 85%. Based on remote sensing information, the species occurs at sites with an annual mean temperature of 19.3 °C, ranging from 11.6 to 26.1 °C during the year. Annual precipitation is about 1843.7 mm with yearly variations from 9.6 to 421.6 mm. Further bioclimatic information is provided in Table 6 .