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Pedesta similissima (Devyatkin, 2002) syn. n. (Fig. 26) Thoressa similissima Devyatkin, 2002: 127, type locality: Thanh Hoa Province, northern Vietnam. = Pedesta submacula (Leech, 1890). Note. Re-examination of the type specimens of Pedesta similissima (Fig. 26, Fig. 27) shows that both the external and genitalic characters are identical to those of P. submacula (Figs. 9���17, Fig. 18), suggesting the two species are conspecific. The analysis of COI gene sequences confirmed this point (Fig. 1). Devyatkin (2002) mistook P. rubella for P. submacula and thus resulted in this synonym. Distribution. Based upon the specimens examined, data from literature (e.g. Ikeda et al. 2001; Wang & Tang 2012; Chen 2016; Zhu et al. 2017) and results of the present study, a distribution map of Pedesta submacula and P. rubella is provided (Fig. 28). It shows that P. submacula is widely distributed from northwestern Qinling Mountains through the southern mainland China to central Vietnam, but in southern Guangxi and northern Vietnam, it is replaced by P. rubella. Both species have not been found sympatrically so far. Although P. submacula is separated by P. rubella into two groups, specimens from China and Vietnam are not distinguishable by stable external or genitalic characters. Therefore, the Vietnamese population of P. submacula is not considered a separate subspecies. Geographical gaps between populations of species caused by natural factors are not rare in the butterfly fauna of Indochina. Disjunctions observed differ in their distance and direction. The majority of disjunctions in the Sino- Himalayan ranges has been observed between populations of northern Vietnam (e.g. Hoang Lien Son mountains) and northern part of central Vietnam (northern Truong Son ridge), and populations of W. China and E. Himalayas. During the last two decades, detailed studies of the Vietnamese butterfly fauna have showed disjunctions in representatives of different taxonomical groups, e.g. Pazala Moore, 1888 (Hu et al. 2018, 2019), Devyatkinia Monastyrskii & U��mura, 2016, Lethe H��bner, 1819 (Lang & Monastyrskii 2016), Ypthima H��bner, 1818 (Monastyrskii & Holoway 2013), etc. Sometimes these disjunctions may be filled with the populations of closely related species, showing the characters of vicariance, the existence of refugiums and the speciation processes occurred in this area. Judging from the modern distribution pattern of Pedesta submacula (Fig. 28), it is very possible that the Vietnamese population of this species was separated from the main distribution range in the course of geological and climatic changes, and then, it adapted to new conditions and lost contact with the main populations; this resulted in the formation of a new taxon, viz. P. rubella. Such a distribution pattern may suggest that the range of this species was larger in the past, when the corresponding habitats shifted down the slopes during cooler and drier glacial period. During the warmer eras, the species may become isolated when these habitats receded to higher altitudes. Such cycles of habitat changes may have also led to varying degrees of divergence in some groups of butterflies. At present, this scenario presents a suitable explanation for range disjunctions of many separate montane butterfly taxa in Vietnam., Published as part of Li, Meng, Monastyrkii, Alexander L., Kolesnichenko, Kirill A., Liu, Zihao, Xue, Guoxi, Long, Jifeng & Tang, Huaxing, 2020, Morphological and molecular characters reveal the status of Pedesta rubella (Devyatkin, 1996) stat. n. and P. similissima (Devyatkin, 2002) syn. n. (Lepidoptera, Hesperiidae), pp. 217-231 in Zootaxa 4743 (2) on pages 224-229, DOI: 10.11646/zootaxa.4743.2.5, http://zenodo.org/record/3687821, {"references":["Devyatkin, A. L. (2002) Hesperiidae of Vietnam, 11. New taxa of the subfamily Hesperiinae (Lepidoptera, Hesperiidae). Atalanta, 33 (1 / 2), 127 - 135.","Ikeda, K., Nishimura, M. & Inagaki, H. (2001) Butterflies of Cuc Phuong National Park in northern Vietnam (5). Butterflies, 30, 58 - 66.","Wang, M. & Tang, D. M. (2012) Butterflies of Guangxi Maoershan National Nature Reserve. Guangxi Nationalities Publishing House, Nanning, 194 pp.","Chen, Z. J. (2016) Guizhou Butterfly. Guizhou Science and Technology Publishing House, Guiyang, 557 pp.","Zhu, L. X., Liu, Z. H., Yu, L. & Ou, Y. Y. (2017) Butterfly Fauna of Anhui. Press of University of Science and Technology of China, Hefei, 429 pp.","Hu, S. J., Cotton, A. M., Condamine, F. L., Duan, K., Wang, R. J., Hsu, Y. F., Zhang, X. & Cao, J. (2018) Revision of Pazala Moore, 1888: The Graphium (Pazala) mandarinus (Oberthur, 1879) Group, with treatments of known taxa and descriptions of new species and new subspecies (Lepidoptera: Papilionidae). Zootaxa, 4441 (3), 401 - 446. https: // doi. org / 10.11646 / zootaxa. 4441.3.1","Hu, S. J., Condamine, F. L., Monastyrskii, A. L. & Cotton, A. M. (2019) A new species of the Graphium (Pazala) mandarinus Group from Central Vietnam (Lepidoptera: Papilionidae). Zootaxa, 4554 (1), 286 - 300. https: // doi. org / 10.11646 / zootaxa. 4554.1.10","Monastyrskii, A. L. & Uemura, Y. (2016) Unique wood nymphs from China and Vietnam: Devyatkinia singularis gen. et spec. nov. Atalanta, 47 (1 / 2), 131 - 137.","Lang, S. Y. & Monastyrskii, A. L. (2016) Description of two new species of the Lethe manzorum - group (Lepidoptera, Nymphalidae, Satyrinae) from China. Zootaxa, 4103 (5), 453 - 462. https: // doi. org / 10.11646 / zootaxa. 4103.5.3"]}

Pedesta rubella (Devyatkin, 1996) stat. n. (Fig. 2, Figs. 3���8) Thoressa submacula rubella Devyatkin, 1996: 603, type locality: near Moc-Chau, northern Vietnam; Monastyrskii & Devyatkin 2015: 77. Thoressa submacula: Devyatkin 2002: 128, fig. 2 (misidentification); Ikeda et al. 2001: 64, fig. 21 (misidentification). Diagnosis. Based upon specimens examined and field investigation, the wing pattern variability of Pedesta rubella is recognized as follows: on the forewing, subapical spots in spaces R 4 ���R 5 may be wide and strip shaped (Fig. 3, Fig. 25b), or narrow and dot like (Figs. 4���7, Fig. 25a); on the dorsal side, the dot in space R 3 may be prominent (Figs. 3, 7) or nearly vanish (Figs. 4���6); the lower cell spot is sometimes elongated toward wing base (Figs. 4, 5), or equally sized with the upper one (Figs. 3, 7), or even smaller (Fig. 6); spots in spaces M 3 and CuA 1 are wide (Fig. 3) or relatively narrower (Fig. 5). On the ventral side of the hindwing, two basal spots in space Sc+R 1 are well separated (Figs. 4, 6, 7, Fig. 25a) or connected (Figs. 3, 5, Fig. 25b); between the basal and submarginal spots in spaces M 1 ���M 2 there is sometimes a pair of white spots which form the postdiscal series with those in spaces Rs, M 3 and CuA 1 (Figs. 6, 7, Fig. 25b). This variability can be observed among individuals of the same group (Fig. 25), and does not represent seasonal forms. Wing patterns of P. submacula fall into this variability (Figs. 9���17), so the two species can hardly be distinguished without an examination of genitalia. The differences in male genitalia of Pedesta submacula (Fig. 18) and P. rubella (Fig. 19, Fig. 20) are as below: 1. In dorsal view, the uncus of Pedesta submacula is obviously wider (it is narrower in P. rubella), and the two distal branches are pointed (they are slender and blunt pointed in P. rubella), between which there is a Ushaped gap (the gap is narrower or V-shaped in P. rubella). 2. In Pedesta submacula, there is usually a pair of short triangular horns on the dorsal side of the tegumen terminally, and sometimes the horns are rather long and sharply pointed (Fig. 26, Fig. 27). But in P. rubella they are vestigial or absent. 3. In lateral view, the distal branches of both valva are robust and semi-erect in Pedesta submacula, but they are usually slender, longer, and erect in P. rubella. Among the specimens of Pedesta rubella, distal branches of valva in some individuals are robust (Fig. 21). Nevertheless, the shape of the uncus is always a good character to distinguish the two species. Note. The holotype of Pedesta submacula is not examined in this study, but the possibility that it belongs to P. rubella can be excluded due to the great distance between the type locality of P. submacula, viz. Changyang (Hubei Province, China) and the range of P. rubella (Fig. 28). Although Pedesta rubella is proved to be a separate species, the unique wing pattern and the remarkable bifurcate distal branch of the right valva of the holotype are conspicuously different from those of the specimens from other localities. If more materials bearing the same morphological characters as the holotype can be found in the future, it will be possible to consider the population from other localities as a distinguishable subspecies of P. rubella. But for now, it is more reasonable to treat the holotype as an individual variation of the species. Description. Females of Pedesta rubella (Fig. 8) and P. submacula (Figs. 11, 13, 15, 17) exhibit similar wing patterns. Unlike the corresponding males, in the lower half of space CuA 2 there is a small triangular patch located near vein 2A, and the line connecting the patch and cell spots is perpendicular to the costa. Genitalia are described as follows to distinguish the females of the two species. Female genitalia of Pedesta rubella (Fig. 22). Papillae anales flabellate in lateral view, covered with long hairs. Apophyses posteriores three times as long as papillae anales. Lamella postvaginalis and lamella antevaginalis well developed and sclerotized. In ventral view, middle of posterior edge of lamella postvaginalis extended to be a fishtail plate, of which basal half tapered and distal margin shallowly concave. Lamella antevaginalis almost triangular, each side of basal area bear an elongated plate with irregular distal margin; middle of posterior edge protruding, round. Ductus bursae tube-like; bursa copulatrix bursiform, constricted in middle, without signum. Female genitalia of Pedesta submacula (Fig. 23). Similar to P. rubella, but extended plate of lamella postvaginalis more developed, shovel-shaped; lamella antevaginalis triangular, top angle blunt, base angle on each side much smaller and sharply pointed. Bionomics. Most specimens of GX-group were caught in limestone karst habitat in Nonggang NNR (Fig. 24). In this subtropical area, Pedesta rubella is common from April to October. The adults fly rapidly and prefer to suck mud or liquid on the ground (Fig. 25). The males also like to occupy the crown of shrubs as a territory and drive out intrusive butterflies. According to our investigations, females of this species appear to be remarkably less numerous than males., Published as part of Li, Meng, Monastyrkii, Alexander L., Kolesnichenko, Kirill A., Liu, Zihao, Xue, Guoxi, Long, Jifeng & Tang, Huaxing, 2020, Morphological and molecular characters reveal the status of Pedesta rubella (Devyatkin, 1996) stat. n. and P. similissima (Devyatkin, 2002) syn. n. (Lepidoptera, Hesperiidae), pp. 217-231 in Zootaxa 4743 (2) on pages 221-223, DOI: 10.11646/zootaxa.4743.2.5, http://zenodo.org/record/3687821, {"references":["Devyatkin, A. L. (1996) New Hesperiidae from north Vietnam, with the description of a new genus (Lepidoptera, Rhopalocera). Atalanta, 27 (3 / 4), 595 - 604.","Monastyrskii, A. L. & Devyatkin, A. L. (2015) Butterflies of Vietnam (an illustrated checklist). 2 nd Edition. Planorama Meida Co., Ltd, Hanoi, 95 pp., 17 pls.","Devyatkin, A. L. (2002) Hesperiidae of Vietnam, 11. New taxa of the subfamily Hesperiinae (Lepidoptera, Hesperiidae). Atalanta, 33 (1 / 2), 127 - 135.","Ikeda, K., Nishimura, M. & Inagaki, H. (2001) Butterflies of Cuc Phuong National Park in northern Vietnam (5). Butterflies, 30, 58 - 66."]}

Mo, Yunming, Zhang, Wei, Zhou, Shichu, Chen, Tianbo, Tang, Huaxing, Meng, Yuanjun, Chen, Weicai (2013): A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China. Zootaxa 3710 (2): 165-178, DOI: http://dx.doi.org/10.11646/zootaxa.3710.2.3

Kaloula nonggangensis sp. nov. Holotype. NHMG 1106036, adult male, from the Nonggang National Nature Reserve, Southern Guangxi Province, China (22.4522 �� N, 106.9354 �� E; altitude: 186 m a.s.l.), collected on June 28, 2011 by Weicai Chen, Yunming Mo (Figure 2 A, B, C). Paratypes. NHMG 1106030 ��� 35, NHMG 1106037 ��� 41, adult males collected at the same place of the locality by Weicai Chen and Yunming Mo on June 29, 2011. NHMG 1108035, adult female, NHMG 1108036 ��� 41 adult males, collected at the near locality of the holotype (22.4809 �� N, 106.9017 �� E; altitude: 167 m a.s.l.) by Yunming Mo, Wei Liao and Zhuqiu Song on August 11, 2011. Diagnosis. Assigned to the genus Kaloula on the basis of the following: medium size (41.4���52.7 mm in 18 adult male, 52.2 mm in one adult female), smooth or slightly rough olive dorsum with irregular dark-green marks and brown spots (Figure 2 A, B); tips of the fingers dilate and truncated (Figure 3 A); nearly full webbing on toes in males and reduced webbing in females (Figure 3 B); two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (Figure 3 C); chest beige with small lemon-colored spots in male (Figure 2 C); ventral epidermal adhesive gland occupies chest and venter; larvae lacking a keratinized jaw sheath and labial teeth. K. nonggangensis sp. nov. is distinguished from it congeners by a combination of (1) medium size (SVL ranging 41.4���52.7 mm in males), (2) smooth or slightly rough olive dorsum with irregular dark-green marks and brown spots, (3) tips of the fingers dilate and truncated, (4) chest beige with small lemon-colored spots in male, and (5) male with two side protuberant osseous tubercles on the upper surface of the tips of fingers. Description of holotype. Adult male; SVL 47.5 mm; habitus rotund, body rounded; head proportionally small, length 25 % of SVL, 76 % of head width; snout slightly pointed beyond lower jaw, its tip round in dorsal aspect and in lateral aspect; eyes protruding laterally beyond silhouette of head in dorsal aspect, protruding very markedly beyond dorsal surface of head in lateral aspect; pupil circular; interorbital region flat with some small tubercles; snout less than half of the head length; upper eyelid width slightly smaller than interorbital distance, and much smaller than snout length; eye diameter (4.63 mm) almost equal to snout length (4.64 mm), longer than interorbital distance (3.8 mm) and eye-narial distance (2.1 mm); canthus rostralis indistinct; loreal region sloping, moderately concave; nostrils barely protuberant, very close to tip of snout (eye-nostril distance longer than distance from nostril to tip of snout); internarial region slightly concave; tympanum hidden and indistinct; tympanic fold prominent, extending from posterior corner of eye to supra-axillary region; tongue oval without posterior notch; choanae elongate oval and small, situated at anterolateral edge of palate, separated by a distance about two times of their diameter; dentigerous process of vomer in posterior edge of choanae evident, almost touching in the middle line but vomerine teeth apparently absent; vocal slits large, just posterior to rictus. Forelimbs. Arms short, forearms not hypertrophied; hand relatively large (HAL 14.3 mm); forearm and hand length (LAHL 23.3 mm) almost as long as half of SVL (47.5 mm); fingers disks obviously dilated and tips truncated; relative length of fingers I Hindlimbs. Hind limbs relatively short and brawny; heels not overlapping for long distance when legs are folded at right angle to body, and when appressed to body, tibiotarsal articulation reaching the shoulder; tibia 2.7 times longer (TL 19.7 mm) than wide (TW 7.4 mm), and shorter than foot length (FOL 21.3 mm), about 42 % of SVL. Relative length of toes: I Skin. Skin of dorsal surfaces of body, head, and limbs smooth; ventral surfaces of throat, trunk, and limbs smooth; loose skin overlying median subgular vocal sac forms slight sternal fold; ventral epidermal adhesive gland occupies sternal region and venter. Color of holotype in life. Dorsal parts of head and dorsum, flank, forelimb, thigh, tibia and foot light olivegreen with dark moss-green marbling; loreal region, tympanic region and tympanum slightly dark olive-green; dorsal parts of two sides shoulder light olive-green; lower part of flanks with some lemon spots; lower jaw, throat, margin of throat and chest beige with some small lemon spots; belly, ventral part of forelimbs and hind limbs creamy white. Color of holotype in preservative. Dorsum brown with dark marbling. Ventral surface of chest, throat, belly, interior portions of arms and thighs cream color. Measurements. Holotype: SVL 47.5, HL 11.7, HW 15.2, SL 4.6, IND 2.8, IOD 4.1, UEW 2.7, ED 4.6, LAHL 23.3, HAL 14.3, HLL 62.7, TL 19.7, TW 7.1, FOL 21.3. Etymology. The species epithet is a Latin adjective referring to the locality in which the new species was collected. The suggested English name is the Nonggang narrow-mouthed frog. Tadpole. Six tadpoles (Voucher No.: NHMG_T 20110801, NHMG_T 20120901 ���05) from stages 36 to 42 sensu Gosner (1960) were collected from the type locality of K. nonggangensis sp. nov. Morphological measurements and characteristics were examined following Altig & McDiarmid (1999). The 16 S rRNA sequence (~ 540 bp) of a specimen (Voucher No.: NHMG_T 20120901) is 99 % identical to a reference sequence of paratype (Voucher No.: NHMG 1108036). Measurements of one tadpole at developmental stage 38: total length 37.5 mm, head-body length 0.54 time of tail length; eyes lateral, small, and visible from ventral view. In dorsal view, body nearly rectangle, snout broadly rounded. Interpupilar distance 0.84 maximum body width, internarial distance 0.17 of interpupilar distance. Width of oral apparatus 0.27 head-body length; lips not expanded; lacking labial teeth, horny beak and papillae. Nostrils closed at stage 38. Spiracle median, opening slightly anterior the end of body, free portion with an arched membrane, the inner wall attached to body wall; anal tube median and elongate, and the inner wall fused to ventral fin. The base of tail musculature strong, its height 0.35 of tail height; maximal tail height located at the proximal 1 / 2 of the tail, caudal muscles tapering gradually. Dorsal fin originates at the tail bodyjunction and the ventral fin originates at the ventral terminus of the body. The dorsal fin is nearly equal to the ventral fin in height. Dorsal fin and ventral fin have some yolk-yellow spots. Especially, dorsal fin near the tail body-junction has a yolk-yellow line. Tail tip broadly rounded. Head and body brownish with some faint yellow pigments (Figure 2 G, H). Variation. Individuals of the type series are generally similar in morphology. The female (NHMG 1108035) is two-third webbed (webbing formula, I 2 - ��� 2 + II 2 - ��� 3 III 2 ��� 3 IV 3 + ��� 2 - V). Two specimens (NHMG 1108035 ��� 36) have some small spinous tubercles in the anal region. Five specimens (NHMG1106033, 1108035���36, 1108038, 1108040) have two subarticular tubercles on the third toe, and the remaining specimens have three subarticular tubercles on the third toe. Four individuals (NHMG 1106040 ���42, 1108037) have a slightly rough dorsum scattered with a few small rounded tubercles. The tympanic fold is indistinct in six individuals (NHMG1106036, 1108037��� 39, 1108040��� 41). All males have two side protuberant osseous tubercles on the upper surface of the tips of fingers. Between two side protuberant osseous tubercles, four individuals have small tubercles ranging from 1 to 4 (NHMG 1106031: L II (left finger II), 3; NHMG 1106033: L II, 4; NHMG 1108036: R I (right finger I), 2; NHMG 1108039: L I, 1; R III, 1), which size is about 1 / 5 of two side protuberant osseous tubercles. Female also have two side protuberant osseous tubercles but which size is only 1 / 2 that of the male. In preservative, seven specimens (NHMG 1106031 ���34, 1106035, 1106039, 1108040) have a gray dorsum, and the remaining specimens are dark brown. Male secondary sexual characters. Nuptial pad absent; external subgular vocal sac present; throat with some lemon spots; chest and belly with epidermal adhesive gland. Ecology. K. nonggangensis sp. nov. were observed in primary or secondary karst evergreen forest, and cultivated fields near the forest. Presently, K. nonggangensis sp. nov. is only known from the Nonggang National Nature Reserve. Sites where we observed the species ranged from 150���200 m elevation. On June 29 and August 11, 2011, the species was observed to gather in the temporary plash after rainstorm. Sequence divergence. Uncorrected sequence divergences between K. nonggangensis sp. nov. sequences and all homologous sequences available on the genus Kaloula are listed in Table 2. The uncorrected p -distance genetic distance between K. nonggangensis sp. nov. and K. verrucosa tissues examined was 1.4���1.6 %, while that between K. nonggangensis sp. nov. and other species was> 1.8 % (Table 2). Preliminary hypothesis of phylogenetic relationship. Among 15 Kaloula species (Frost 2013), only 9 species' sequences (including K. baleata, K. borealis, K. conjuncta, K. mediolineata, K. picta, K. pulchra, K. rugifera, K. taprobanica, K. verrucosa) are available from GenBank. Based upon our preliminary molecular data, the genus Kaloula formed a monophyletic group with well-supported values (BP= 98; BBP=1.00), and was divided three clades (Figure 4). Clade A consists of Kaloula taprobanica Parker, 1934; Clade B consists of K. borealis, K. rugifera, K. verrucoca, and K. nonggangensis sp. nov. and Clade C consists of K. baleata van & Muller, 1836, K. conjuncta Peters, 1863, K. mediolineata Smith, 1917, K. picta Dum��ril & Bibron, 1841, and K. pulchra Gray, 1831. Comparisons: K. nonggangensis sp. nov. differs from all other species of Kaloula by having smooth or slightly rough dorsum without rough tubercles, finger tips widely expanded and truncated, two side protuberant osseous tubercles on the upper surface of the tips of fingers in male, throat and chest with some small lemon spots in male, and larvae with some yolk-yellow spots in body and tail. K. nonggangensis sp. nov. is most similar in appearance to K. borealis, K. rugifera and K. verrucosa (Figure 2). However, K. nonggangensis sp. nov. differs from K. borealis by having widely expanded terminal digital disks (vs. lacking expanded terminal digital disks in the latter); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (without osseous tubercles in the latter, Figure 3 D); by having nearly full webbing (vs. 1 / 2 webbing in the latter); by throat and chest beige with some small lemon spots (vs. distinct black spots in the throat area in the latter) (Fei et al. 2009). Furthermore, in life, the tadpole of K. borealis has dark brown dorsal body and tail, without pigment and venter white or hoar (vs. the tadpole of K. nonggangensis sp. nov. has brownish head and body with some faint yellow pigments and dorsal fin near the tail body-junction has a yolk-yellow line) (Fei et al. 2009; Zhou et al. 2011). K. nonggangensis sp. nov. differs from K. rugifera by having a relatively bigger body size (SVL 41.4���52.7 mm in male vs. 35.5 ���43.0 mm in male for K. rugifera) (Table 3). It can be further distinguished from K. rugifera by finger tips widely expanded and truncated (slightly expanded in K. rugifera, Figure 3 F); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (having two clusters of osseous tubercles instead of two side protuberant osseous tubercles in the latter) (Figure 3 F); by throat and chest with some small lemon spots in male (vs. cream without pigments); by larvae with some yolk-yellow spots in body and tail (vs. dorsum and tail dark brown, no pigment) (Fei et al. 2009; Zhou et al. 2011). K. nonggangensis sp. nov. also differs from K. verrucosa by smooth or slightly rough dorsum without rough tubercles (rough dorsal skin and tubercles present in K. verrucosa) (Figure 2 F); by finger tips widely expanded and truncated (non-expansion of terminal digital disks in the latter, Figure 3 E); by two side protuberant osseous tubercles on the upper surface of the tips of fingers in male (4���6 free osseous tubercles in the latter) (Figure 3 E); by larvae with some yolk-yellow spots in body and tail (vs. dorsum and tail dark brown with black pigment in the latter) (Fei et al. 2009). The morphological differences in the shape of terminal disks (Figure 3) have been assessed in large numbers of specimens in the studies of 83 for K. borealis, 100 for K. verrucosa and 67 for K. rugifera, partly in specimens from different localities (Kunming and Dali in K. verrucosa) (Appendix I, Fei et al. 2009). K. nonggangensis sp. nov. differs from K. aureata Nutphand, 1989, by having an olive dorsum without lateral bands (vs. bright yellow in median dorsum, and yellowish orange lateral bands along the back) (Pauwels & Ch��rot, 2006). K. nonggangensis sp. nov. differs from K. baleata by having a ventral adhesive gland in males (vs. venter without epidermal adhesive gland); by having olive dorsum without rough tubercles (vs. brownish dorsum with rough tubercles, and a bright yellow spots near axilla in the latter) (Pauwels et al. 1999). K. nonggangensis sp. nov. differs from K. conjuncta, K. kalingensis, K. kokacii and K. walteri by a larger body size (SVL 41.4���52.7 mm in males vs. 26.6���30.1 mm in K. conjuncta, 24.2���32.9 mm in K. kalingensis, 37.0��� 39.1 mm in K. kokacii, 24.5���31.5 mm in K. walteri (Diesmos et al. 2002). It also differs from K. conjuncta by the absence of rough dorsal tubercles (vs. tubercles distributed over the entire dorsum); by having olive dorsum (vs. brownish dorsum) (Diesmos et al. 2002). It also differs from K. kalingensis and, K. kokacii by its ventral adhesive gland in males (vs. venter without epidermal adhesive gland) (Brown et al. 2000; Diesmos et al. 2002). It also differs from K. walteri by the presence of a round, small, and pointed outer metatarsal tubercle (vs. absence, or presence of only a faint, very reduced outer metatarsal tubercle); by the presence of three subarticular tubercles on the fourth toe (vs. two); by having widely expanded terminal digital disks (vs. lacking expanded terminal digital disks in the latter) (Diesmos et al. 2002). K. nonggangensis sp. nov. clearly differs from K. assamensis, K. mediolineata, K. pulchra and K. taprobanica by absence (vs. presence) of dorsolateral bands. It also differs from K. assamensis by its inner metatarsal tubercle smaller (vs. larger) than first toe; lacking a vertebral stripe (vs. presence of a dark edged lemon vertebral stripe) (Das et al. 2004; Nath et al. 2011). It also differs from K. mediolineata by its finger tips widely expanded (vs. non expanded) and truncated (Diesmos et al. 2002). It also differs from K. pulchra by its smaller body size (SVL 41.4��� 52.7 mm vs. 55.0���77.0 mm in K. pulchra (Fei et al. 2009); snout length about 1 / 3 head length (vs. half of the head length; by smooth venter (vs. granular venter) (Fei et al. 2009). It can be further distinguished from K. taprobanica by lacking (vs. having) large irregular shaped markings of yellow-red color in dorsum median area (Dutta & Manamendra-Arachchi 1996). K. nonggangensis sp. nov. differs from K. picta and K. rigida by absence (vs. presence) of supernumerary tubercles at the base of each digit of the manus. It also differs from K. picta by its inner metatarsal tubercle shorter (vs. equal to or longer) than first toe (Diesmos et al. 2002). Finally, K. macrocephala (originally treated as a synonym of K. pulchra) differs from K. nonggangensis sp. nov. by having indistinct dorsolateral bands and mid-dorsum covered by large-sized irregular patches (vs. clearly absent dorsolateral bands in new species) (Bourret 1942; Ohler 2003; Pauwels & Ch��rot 2006). In addition, genetically, except for K. borealis, K. rugifera and K. verrucosa, the uncorrected sequence divergences between K. nonggangensis sp. nov. 16 S rRNA sequences and all homologous sequences available on GenBank (Table 2) were greater than 3 %, a value usually representing differentiation at the species level in frogs (Vences et al. 2005; Fouquet et al. 2007). Discussion Fei et al. (2009) divided Chinese Kaloula species into two groups: the K. pulchra group (consisting of K. pulchra) and the K. verrucosa group (consisting of K. borealis, K. rugifera and K. verrucosa), being consistent with our phylogenetic trees. K. nonggangensis sp. nov. embedded within the K. verrucosa group. K. nonggangensis sp. nov., K. borealis, K. rugifera and K. verrucosa formed a monophyletic group with high supported values (Figure 4). Preliminary molecular data indicated that K. nonggangensis sp. nov. and K. borealis were sister species. However, due to weak inter-nodes support values (K. verrucosa group are lower than the values usually representing differentiation at the species level in frogs (Vences et al. 2005; Fouquet et al. 2007) (Table 2). According to the results of these authors, in most cases species are differentiated by uncorrected p -distances in 16 S rRNA of 3 % or higher. If the criterion was strictly applied in this group, then K. nonggangensis sp. nov., K. borealis, K. ru, Published as part of Mo, Yunming, Zhang, Wei, Zhou, Shichu, Chen, Tianbo, Tang, Huaxing, Meng, Yuanjun & Chen, Weicai, 2013, A new species of Kaloula (Amphibia: Anura: Microhylidae) from southern Guangxi, China, pp. 165-178 in Zootaxa 3710 (2) on pages 168-175, DOI: 10.11646/zootaxa.3710.2.3, http://zenodo.org/record/216543