Amphibia: Anura: Megophryidae

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Article. 541 https://doi.org/10.11646/zootaxa.4324.3.8 .... China: T aoyuandong N ature R eserve, H unan. SY. S a002123. MF667904. K. J560396 ...... ralesp in o sa. 36.2. -38.0. /. -. ++. +. -. ++. +++ sm all. 0.55. X. pa rva. 37.0. -44.0. 45.0. -54.0.
Zootaxa 4324 (3): 541–556 http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

https://doi.org/10.11646/zootaxa.4324.3.8 http://zoobank.org/urn:lsid:zoobank.org:pub:C4D32E0E-7330-4030-9399-70ED07D24EAD

A new species of the genus Xenophrys (Amphibia: Anura: Megophryidae) from an offshore island in Guangdong Province, southeastern China JIAN WANG1, ZU-YAO LIU1, ZHI-TONG LYU1, ZHAO-CHI ZENG1 & YING-YONG WANG1, 2 1

State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275, P.R. China. 2 Corresponding author. E-mail: [email protected]

Abstract A new species, Xenophrys insularis sp. nov., is described based on a series of specimens collected from Nan’ao Island, Shantou City, Guangdong Province, China. The new species is a separately evolving lineage that was robustly supported by the phylogenetic analysis based on the mitochondrial CO1 + 12S rRNA + 16S rRNA genes; morphologically, it can be distinguished from other known congeners by the following combination of characteristics: body size moderate, SVL 38.6–41.2 mm in adult males, 47.1 mm in a single adult female; snout obtusely rounded in dorsal view; head width slightly larger than head length; strong vomerine ridge bearing vomerine teeth; margin of tongue weakly notched behind; supratympanic fold forming a large swollen supraaxillary gland above insertion of arm; hindlimbs relatively short, heels not meeting when thighs are held at right angles to body, tibio-tarsal articulation reaches forward the posterior edge of tympanum; fingers and toes without lateral fringes, with rudiment of web at bases of toes, distinct subarticular tubercle only at the base of each finger and toe; single vocal sac in males; gravid female bears pure white oocytes. Xenophrys insularis sp. nov. is endemic to the Nan’ao Island and represents the first Xenophrys species found on and possibly endemic to a small offshore island. Key words: Xenophrys insularis sp. nov., mitochondrial DNA, morphology, taxonomy, insular endemic species

Introduction The Asian horned toad, genus Megophrys sensu lato Kuhl & Van Hasselt of subfamily Megophryinae in the family Megophryidae, is characterized by the horn-like tubercles on the edge of the eyelids in most of congeners, comprising 61 species dwelling in montane forest area in subtropical and tropical Asia. For decades, these toads has suffered an intensive taxonomic debate. In a recent publication, based on a large-scale molecular analysis on megophryine species, Chen et al. (2017) considered that Megophrys sensu lato is divided into three previously established genera namely Megophrys sensu stricto, Xenophrys Günther and Atympanophrys Tian & Hu. Subsequently, Mahony et al. (2017) provided a similar phylogenetic result based on a larger molecular dataset and partial morphological evidences, proposing to regard the entire subfamily Megophryinae as a single genus, Megophrys, and to treat seven previous genera, Megophrys, Xenophrys, Panophrys Rao & Yang, Atympanophrys, Ophryophryne Boulenger, Pelobatrachus Beddard and Brachytarsophrys Tian & Hu, as subgenera within the genus Megophrys. Comparing these two classification opinions, we rather follow Chen's taxonomy because Megophrys, Atympanophrys, Ophryophryne and Brachytarsophrys are recognized genera (Chen et al. 2017; Fei et al. 2012) which fulfill the following criteria: being monophyletic, reasonably compact, and morphologically distinct at least (Gill et al. 2005). Currently, the genus Xenophrys contains 48 species, distributed in the southern and eastern Himalayas, Indochina Peninsula, and northward to the Qinling Mountains and Huangshan Mountains of mainland China (Chen et al. 2017; Wang et al. 2014; Wang et al. 2017). A total number of 33 recognized species of the genus Xenophrys occur in China, of which only 11 species were recorded from eastern and southern China, namely X. mangshanensis (Fei & Ye) distributed widely from Nanling Mountains and Jiulianshan Mountains, X. brachykolos

Accepted by M. Vences: 21 Aug. 2017; published: 28 Sept. 2017

541

(Inger & Romer) from Hong Kong, X. boettgeri (Boulenger) and X. kuatunensis (Pope) from Wuyi Mountains, X. huangshanensis (Fei & Ye) from Huangshan Mountains, and X. jinggangensis Wang, X. lini Wang & Yang and X. cheni Wang & Liu from Mount Jinggang region, situated at the border between the Jiangxi and Hunan Provinces; X. acuta (Wang, Li & Jin) and X. obesa (Wang, Li & Zhao) from Heishiding Natural Reserve, situated in the northern region of the Yunkai Mountains, western Guangdong Province; and X. lishuiensis Wang, Liu & Jiang, from Lishui City, Zhejiang Province. Nan’ao Island (23°11'–23°32' N, 116°53'–117°19' E) (Figure 1), Shantou City, Guangdong Province, China, as a small offshore island, covers an area of only 128 km2, away from the mainland at a nearest distance of 7 km, where the Tropic of Cancer crossing the center core area. During our repeated herpetological surveys from 2008 to 2015 in this offshore island we collected a series of specimens of genus Xenophrys, which have been reported as Megophrys minor Stejneger (now Xenophrys minor) by Li et al. (2011). However, our phylogenetic analyses and morphological comparisons showed that the population from Nan’ao Island significantly differs from X. minor and all other recognized Xenophrys species, and represents an undescribed taxon. Therefore, we here describe this population as a new species.

FIGURE 1. Sampling localities of Xenophrys toads in this study. 1: Nan’ao Island, Guangdong; 2: Hong Kong; 3: Heishiding Nature Reserve, Guangdong; 4: Mt. Longtou, Guangdong; 5: Nanfengmian Nature Reserve, Hunan; 6: Mt. Jinggang, Jiangxi; 7: Taoyuandong Nature Reserve, Hunan; 8: Guadun Village (=Kuatun), Fujian; 9: Mt. huangshan, Anhui; 10: Dujiangyan City, Sichuan.

Material and methods Sampling. For molecular analyses, we selected a total of 37 samples (muscle tissues) from all 12 species of the genus Xenophrys recorded in southern and eastern China, and of the undescribed taxon from Nan’ao Island, Guangdong, China. Except three samples of X. mangshanensis, samples were taken from topotypical specimens of X. brachykolos, X. boettgeri, X. kuatunensis, X. acuta, X. obesa, X. jinggangensis, X. lini, X. cheni, X. huangshanensis, X. minor; all collection details are listed in Table 1, and marked in Figure 1. Besides, two sequences of the genus Leptolalax downloaded from Genbank were used as the out group (see Table 1).

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650 retained) and numbers before slash indicate Bayesian posterior probabilities (>0.70 retained).The symbol “-” represents bootstrap value below 60.

Xenophrys insularis sp. nov. Figure 3 & 4. Chresonymy: Megophrys minor Stejneger, 1926: Li et al. 2011.

Holotype. SYS a002169, adult male (Figure 3: A to D, Figure 4), collected on 29 June 2013 by Yu-Long Li (YLL hereafter), Jian Zhao (JZ hereafter) and Run-Lin Li (RLL hereafter) from Nan’ao Island (23°26'0.09" N, 117°4'45.61" E; 425 m a.s.l.), Shantou City, Guangdong Province, China (Figure 1) . Paratypes. Five specimens (four adult males, one adult female), from Nan’ao Island at elevations between 50– 500 m a.s.l., SYSa002168 (Figure 3: E and F), adult female, SYS a002167, 2170–71, adult males, collected on 29 June 2013 by YLL, JZ, and RLL, SYS a003666 / CIB 106881, adult male, collected on 29 April 2015 by Jian Wang (JW hereafter) and Can-Rong Lin (CRL hereafter). Diagnosis. Xenophrys insularis sp. nov. is characterized by the combination of following characters: (1) Body stout and moderate-sized, SVL 36.8–41.2 mm in four adult males, 47.1 mm in the single adult female specimen; (2) head width slightly larger than head length; (3) snout obtusely rounded in dorsal view; (4) tympanum distinct and small, TD/ED ratio 0.46–0.57; (5) vomerine ridge strong with vomerine teeth; (6) margin of tongue weakly notched behind; (7) hindlimbs short, heels not meeting, tibio-tarsal articulation reaches the posterior edge of A NEW SPECIES OF XENOPHRYS FROM CHINA

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tympanum on each side; (8) fingers and toes without lateral fringes; (9) toes with rudiment of webs at their bases; (10) subarticular tubercle only present at the base of each finger and toe; (11) dorsal skin rough with several large warts scattered on flanks, usually short skin ridges occurred on dorsal body; (12) presence of a barely visible hornlike tubercle at the edge of the eyelid; (13) supratympanic fold narrow, forming a swollen supraaxillary gland above insertion of arm; (14) greyish brown or dark olive dorsally, dorsal coloration uniform with only a barely visible incomplete dark triangle present between eyes and several dark brown cross-bars on dorsal limbs; (15) presence of a single vocal sac in males; (16) presence of two darker brown nuptial spines on dorsal surface of the first and second fingers during breeding season, respectively. Comparisons. Comparative data of Xenophrys insularis sp. nov. with 48 recognized species of the genus Xenophrys were obtained from the literature (Boulenger 1908; Inger & Romer 1961; Taylor 1962; Fei et al. 1992; Tian et al. 2000; Ohler et al. 2002; Mathew & Sen 2007; Fei et al. 2009; Mahony et al. 2010; Mahony 2011; Mahony et al. 2011; Stejneger 1926; Wang et al. 2012; Mahony et al. 2013; Li et al. 2014; Wang et al. 2014; Orlov et al. 2015; Wang et al. 2017) or from own examination of individuals of X. acuta, X. boettgeri, X. brachykolos, X. cheni, X. jinggangensis, X. kuatunensis, X. mangshanensis, X. maosonensis (Bourret), X. minor, X. lini and X. obesa, as listed in Appendix 1. All comparative morphological data are shown in Tables 3 and 4. Xenophrys insularis sp. nov., was previously reported as X. minor (Li et al. 2011), and differs from true X. minor from type locality by having obtusely rounded snout (vs. pointed snout); presence of the horn-like tubercles above the eye (vs. absent); comparative tympanum size, TD/ED ratio 0.46–0.57 (vs. enlarged tympanum, TD/ED ratio 0.8–0.83), tympanum diameter being almost equidistant with tympanum–eye distance, TED/TD ratio 0.92– 1.00 (vs. TD greater than TED, TED/TD ratio 0.61–0.62); shank short, TIB/SVL ratio 0.38–0.43 (vs. TIB/SVL ratio 0.46–0.48); tibio-tarsal articulation reaching the posterior edge of tympanum (vs. reaching the region between eye and tip of snout); the heels not meeting when thighs are positioned at right angles to the body in the new species (vs. heels meeting or overlapping). Xenohprys insularis sp. nov. significantly differs from ten species occurring in eastern and southern China by the following combination of characters: presence of vomerine teeth (vs. absence in X. brachykolos, X. boettgeri, X. kuatunensis, X. huangshanensis, X. lini, X. cheni, X. acuta, X. obesa, X. lishuiensis); margin of tongue notched behind (vs. not notched in X. brachykolos, X. jinggangensis, X.lini, X. acuta, X. obesa, X. lishiuensis); toes without lateral fringes (vs. with lateral fringes in X. boettgeri, X. kuatunensis, X. jinggangensis, X. lini, X. cheni, X. acuta); hindlimbs comparatively short, with heels not meeting when thighs are positioned at right angles with respect to body (vs. hindlimbs comparatively longer, with heels meeting or overlapping in X. boettgeri, X. kuatunensis, X. jinggangensis, X. lini, X. cheni). Body size of the new species is smaller than that of the following 20, additional species of Xenophrys, with no overlap of values: X. aceras (Boulenger), X. auralensis (Ohler, Swan & Daltry), X. binglingensis (Jiang, Fei & Ye), X. caudoprocta (Shen) , X. damrei (Mahony), X. glandulosa (Fei, Ye & Huang) , X. jingdongensis (Fei & Ye), X. lekaguli (Stuart, Chuaynkern, Chan-ard & Inger) X. longipes (Boulenger), X. major (Boulenger), X. mangshanensis, X. maosonensis, X. medogensis (Fei, Ye & Huang) , X. megacephala (Mahony, Sengupta, Kamei & Biju), X. omeimontis (Liu), X. robusta (Boulenger) , X. sangzhiensis (Jiang, Ye & Fei), X. shuichengensis (Tian & Sun), X. spinata (Liu & Hu), X. takensis (Mahony). Body size of the new species is larger than in the following eight additional species with no overlap values: X. acuta, X. cheni, X. kuatunensis, X. lishuiensis, X. pachyproctus (Huang), X. vegrandis (Mahony, Teeling & Biju), X. wuliangshanensis (Ye & Fei) , X. wushanensis (Ye & Fei) . Comparative data see Table 3. From the remaining 13 species of the genus which share a moderate body size with the new species, X. insularis sp. nov. can be steadily distinguished by the following combination of characters: hindlimbs short, heels not meeting each other when thighs are adpressed at right angles with respect to body (vs. relatively long hindlimbs, heels meeting or overlapping in X. palpebralespinosa (Bourret), X. parva (Boulenger), X. tuberogranulatus (Shen, Mo & Li), X. zhangi (Ye & Fei), X. daweimontis (Rao & Yang), X. binchuanensis (Ye & Fei), X. ancrae (Mahony, Teeling & Biju), X. oropedion (Mahony, Teeling & Biju); tibio-tarsal articulation reaches the posterior edge of tympanum (vs. reaches the anterior region of eye in X. serchhipii, reaches the middle of eye in X. zunhebotoensis Mathew & Sen); vomerine teeth present (vs. absent in X. baolongensis (Ye, Fei & Xie)); margin of tongue notched behind (vs. not notched in X. oropedion, X. parallela (Inger & Iskandar), X. palpebralespinosa, X. parva, X. tuberogranulatus and X. latidactyla (Orlov, Poyarkov & Nguyen)); toes with rudiment of webs (vs. at least one-fourth webbed in X. ancrae and X. serchhipii).

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FIGURE 3. A and B: Dorsolateral and ventral views of the adult male holotype SYS a002169 of Xenophrys insularis sp. nov. in life; C and D: Ventral views of hand and foot of the holotype in life; E and F: Dorsolateral and ventral views of the adult female paratype SYS a002168 in life.

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FIGURE 4. A, B and C: Dorsal, ventral and lateral views of the adult male holotype SYS a002169 of Xenophrys insularis sp. nov. in preservative; D: Front view of head the holotype in preservative; E and F: Ventral views of hand and foot of the holotype in preservative.

Description of holotype. Adult male. Habitus stout, body size moderate, SVL 39.7 mm; head length slightly shorter than head width, HDW/HDL ratio 1.09; snout rounded nearly to truncate in dorsal view, slightly projecting, sloping backward to mouth in profile, protruding beyond margin of lower jaw; top of head flat; eye large, EYE 41% of HDL, pupil vertical; canthus rostralis well developed; loreal region slightly sloping, not concave; internasal distance slightly larger than interorbital distance; tympanum large, rounded, distinctly visible, TMP/EYE ratio 0.57; choanae large, ovoid at the bases of the maxilla; strong vomerine ridges with vomerine teeth; margin of tongue weakly notched behind. Radioulna length 25% of SVL, hands without webbing, hand length 25% of SVL; no lateral fringes, relative finger lengths: II < I ≤ III < IV; tips of digits rounded, slightly dilated; a large subarticular tubercle at the base of each finger; hindlimbs of moderate length, slightly robust; tibio-tarsal articulation reaches forward to the posterior edge of tympanum when hindlimb is stretched along the side of the body; heels not meeting when thighs are held at right angles to the body axis; tibia length 42% of SVL; foot length 60% of SVL; relative toe lengths: I < II < V < III < IV; tips of toes rounded, slightly dilated; toes with rudiment of webs at their bases; subarticular tubercles absent; no lateral fringes; external tarsal fold present; inner metatarsal tubercle long ellipsoid; outer metatarsal tubercle absent. Dorsal skin rough, with numerous granules and tubercles; several large warts scattered on flanks; an indistinct horn-like tubercle at the edge of the eyelid, barely visible in ethanol; supratympanic fold distinct, curving posteroventrally from posterior corner of eye to a level above insertion of arm and forming a raised supraaxillary gland on each side; ventral surface smooth, with small granules on throat and chest, posteriorly densly covered with pustules and small granules on belly, ventral side and rear of thighs; pectoral gland closer to axilla than to midventral line; single femoral gland on rear of thigh.

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X. huangshanensis

X. jingdongensis

X. jinggangensis

X. kuatunensis

X. latidactyla

X. lekaguli

X. lini

X. lishuiensis



X. caudoprocta

±

±

X. brachykolos

±

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±

X. boettgeri



±

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X. binlingensis

X. daweimontis

±



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X. binchuanensis

X. damrei



±

X. baolongensis

±





X. auralensis

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X. ancrae

X. cheni



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X. aceras



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62.5

61.9–83.4

57.2–68.0

45.9–53.4

34.5–41.2

35.6

56.0–59.5

32.8–39.2

35–36

40–48

36.2–38.0

37.0–44.0

/

54.7

37.1

99.8–115.6

47.2–54.4

47.3–53.0

33.2–39.0

27.5–30.6

27.3–31.6

30.4–35.5

32.5–37.2

X. mangshanensis

X. maosonensis

X. medogensis

X. megacephala

X. minor

X. obesa

X. omeimontis

X. oropedion

X. pachyproctus

X. parallela

X. palpebralespinosa

X. parva

X. robusta

X. sangzhiensis

X. serchhipii

X. shuichengensis

X. spinata

X. takensis

X. tuberogranulatus

X. vegrandis

X. wuliangshanensis

X. wushanensis

X. zunhebotoensis

27

32.5–37.2

77.0

X. zhangi

47.0

X. major

males

SVL

X. longipes

TABLE 3. (Continued)

/

/

38.4

41.3

/

50.5

72.9

54.0–55.0

102.0–118.3

/

/

114.0

45.0–54.0

/

/

36

44.1–48.7

68.0–72.5

37.5–41.2

/

64.4

/

/

73.0

94.0

65.0

females

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Tongue: notched (++), feebly notched (+), or not notched (‒)



+

+

Vomerine teeth: present (+), or absent (‒)

+

+

Horn-like tubercle at edge of upper eyelid: long point (+++); slightly large (++), small (+), absent or indistinct (‒)

+



Genetic sequencing: sequenced (+) or not (‒)

+

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Toes: at least one-fourth webbed (+++), at most one-fourth webbed (++), with rudiment of web (+), or without web (‒)

+

lateral fringes on toes: wide (++), narrow (+), lacking (‒)

0.50

0.50

0.50

0.50

0.44–0.56

0.50

0.71

0.43

0.67

0.39

0.62

0.50

0.40

small