Megophrys (Panophrys) kuatunensis Pope, 1929 Figs. 8 & 9; Table 5. Megalophrys kuatunensis Pope 1929:1 (partim: see Remarks section). –– Megophrys kuatunensis Gee & Boring 1929:20. –– Megophrys (Megophrys) kuatunensis Dubois 1980:472. –– Panophrys kuatunensis Rao and Yang 1997:98. –– Xenophrys kuatunensis Delorme et al. 2006:17. –– Megophrys (Panophrys) kuatunensis Mahony et al. 2017:755. Holotype: Adult male (AMNH 30126), type locality: “Kuatun [Village], Chungan Hsien, northwest Fukien Province, China, 5500–6000 feet.” (=Guadun [ca. 27°40’N, 117°40’E, ca. 1675–1830 m asl], Wuyishan County, Nanping Prefecture, Fujian Province), collected by Clifford H. Pope, April–September 1926 (Pope 1929, 1931). Paratypes: AMNH 30123–30124; AMNH 30239–30258, FMNH 24406 (formerly AMNH 30230), FMNH 24408 (formerly AMNH 30232), FMNH 24411–24413 (formerly AMNH 30235–30237), BMNH 1961.956 (formerly AMNH 30238, then FMNH 24414), BMNH 1985.1294 (formerly AMNH 30234, then FMNH 24410), BMNH 1985.1295 (formerly AMNH 30231, then FMNH 24407), MCZ 28297 (formerly AMNH 30233, then FMNH 24409). Examined specimens: Full morphological datasets were taken for five adult males, holotype and paratypes (AMNH 30126, 30240–30241, 30243–30244), and four adult female paratypes (AMNH 30242, BMNH 1961.956, BMNH 1985.1295, FMNH 24411). SVL measurements were taken for three additional adult male paratypes (FMNH 24406, FMNH 24412–24413). Holotype description: (Figs. 8 & 9; Table 5 for measurements): Sexually mature adult male. Head moderately small, width subequal to length; snout rounded in dorsal view, obtusely protruding in lateral view, without rostral appendage (Fig. 8); loreal region vertical and weakly concave; canthus rostralis angular; dorsal region of snout slightly concave; eye diameter greater than three times as long as maximum tympanum diameter, and longer than snout; eye-tympanum distance subequal to maximum tympanum diameter; tympanum circular, its upper edge not concealed by supratympanic ridge (Fig. 8); nostril orientated laterally, closer to eye than snout; internarial distance greater than eyelid width, and subequal to narrowest point between upper eyelids; pineal ocellus not visible externally; vomerine ridges and vomerine teeth absent; tongue moderately large, weakly notched posteriorly, with no medial lingual process. Forelimbs moderately short and thin, forearm moderately enlarged relative to upper forelimb, and shorter than hand; fingers short and narrow without lateral fringes (Fig. 9B), finger length formula IV ”), parietoscapular-middorsal ridge (“>–”), complete parietoscapular-sacral ridge (“>- <“). The holotype is the only male with no dermal asperities. AMNH 30240 has small white spinular dermal asperities sparsely distributed on tops of tubercles and on the parietoscapular fold on the anterior dorsum, becoming moderately dense posteriorly; on AMNH 30241, asperities are visible from approximately mid trunk length, increasingly in density posteriorly; on AMNH 30242 and AMNH 30243, asperities are sparse on the tympanic region and on posterior lateral surfaces of head and anterior lateral dorsum, sparse on anterior central dorsum but from mid dorsum increases in density posteriorly. Asperities are absent from all remaining surfaces on these specimens. FMNH 204411 is the only female with dermal asperities with just a few scattered on the posterior dorsum. Colouration in preservation of examined paratypes differed from holotype based on the following characters: triangular marking between the eyes had a light central spot on AMNH 30243 and AMNH 30244; dark brown dorsolateral stripe is present on AMNH 30244; AMNH 30240, AMNH 30241 and AMNH 30243 have two to three transverse crossbars on shanks; palmar, thenar and inner metatarsal tubercles distinctly lighter than surrounding surfaces on paratypes; dark brown blotches on gular region, chest and abdomen vary in intensity and density from almost absent on AMNH 30240 to mottled on AMNH 30241; pectoral and femoral glands noticeably lighter than surrounding surfaces on all examined paratypes; ventral surfaces of thighs and shanks faintly mottled on AMNH 30240, AMNH 30241 and AMNH 30244. Secondary sexual characters: Males: weakly raised nuptial pads present, appearing smooth and translucent, or covered with brown microspinules, covering most of the dorsal surface of Finger I, narrowing distally and extending to the base of the distal phalange; nuptial pad absent on Finger II; vocal sac indistinct in preservation; internal vocal slits present near the rear of the mandible; forearms slightly to moderately enlarged relative to the upper forelimbs. Females: mature ova without pigmented poles; nuptial pads, internal vocal slits and enlarged forearms absent. Distribution (Fig. 10): To our knowledge, the presence of M. kuatunensis has only been positively confirmed with molecular data in Fujian and Jiangxi provinces (Li et al. 2014; Wang et al. 2014; Chen et al. 2017). The IUCN Red List assessment for this species indicates a much wider distribution (Huiqing et al. 2004) including locations in Zhejiang, Hunan and Guangxi provinces. Geological barriers exist between these locations that may limit the dispersal of Megophrys. The identity of populations assigned to M. kuatunensis in Zhejiang, Hunan and Guangxi provinces should be confirmed with molecular data. Remarks: Pope (1929) provided a relatively detailed description of the holotype of his new species, M. kuatunensis, including a brief comparison with M. boettgeri and M. minor. He later expanded on this description providing a figure of the holotype, a detailed variation section, and morphological and biological comparison with the sympatric M. boettgeri (Pope 1931). The type series of Megophrys kuatunensis was based on the holotype and 31 paratypes (Pope 1929). Pope (1931:445) included an additional specimen as a paratype, AMNH 30324, stating that “An additional specimen has been found since the appearance of the original description which listed only 31 paratypes ”. This action does not validate AMNH 30324 as a paratype, and thus this specimen is not available for lectotype designation in the future should the holotype become lost/destroyed (International Commission of Zoological Nomenclature 1999). Within the section titled “Notes on paratypes ”, Pope (1929) includes a statement that “Tadpoles were secured”, neither designating a museum number, nor counting them among the 31 specimens referred to directly as paratypes, indicating that Pope did not intend to regard these tadpoles as paratypes. Pope (1931) further comments that of the two series of tadpoles (AMNH 30606 and AMNH 30645) collected from the type locality, he would only tentatively assign the former to M. kuatunensis. These tadpoles should not be considered to be paratypes of M. kuatunensis. Marx (1958) lists three paratypes in the CNHM (FMNH 24408, 24412, 24413), omitting two paratypes FMNH 24406 (formerly AMNH 30230) and FMNH 24411 (formerly AMNH 30235) from his annotated catalogue of type specimens in the collection. This omission was not amended in supplementary catalogue of FMNH types (Marx 1976), however, we confirm that both of these specimens were present in the collection during a recent visit (S. Mahony, pers. com.). In their description of a new species, M. brachykolos, Inger and Romer (1961) examined part of the type series of M. kuatunensis and were the first to note that the paratype series of M. kuatunensis is heterogeneous in species composition. They identified a M. kuatunensis female paratype FMNH (as CNHM) 24408 as M. brachykolos on the basis of several measurements and characters that fell within the variation range of the female M. brachykolos types. They designated this specimen as a paratype of their new species (Inger and Romer 1961; Marx 1976). Our examination of approximately half of the type series of M. kuatunensis (including FMNH 24408) revealed two additional paratypes (AMNH 30239 and BMNH 1985.1294 [formerly AMNH 30234, then FMNH 24410]) that are morphologically conspecific with FMNH 24408, and not M. kuatunensis sensu stricto. Though we partially agree with Inger and Romer (1961) in so far as these specimens share superficial similarities with M. brachykolos sensu stricto (and its sister taxon M. acuta), we suggest that their identification should be considered tentative pending further study of this population due to geographic distance from the type locality of M. brachykolos, and the recognition of extensive cryptic diversity in Megophrys (Chen et al. 2017; Mahony et al. 2017).

Published as part of Tapley, Benjamin, Cutajar, Timothy, Mahony, Stephen, Nguyen, Chung Thanh, Dau, Vinh Quang, Nguyen, Tao Thien, Luong, Hao Van & Rowley, Jodi J. L., 2017, The Vietnamese population of Megophrys kuatunensis (Amphibia: Megophryidae) represents a new species of Asian horned frog from Vietnam and southern China, pp. 465-492 in Zootaxa 4344 (3) on pages 482-486, DOI: 10.11646/zootaxa.4344.3.3, http://zenodo.org/record/1043687