Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences

The question whether taxonomic descriptions naming new animal species without type specimen(s) deposited in collections should be accepted for publication by scientific journals and allowed by the Code has already been discussed in Zootaxa (Dubois & Nemésio 2007; Donegan 2008, 2009; Nemésio 2009a–b; Dubois 2009; Gentile & Snell 2009; Minelli 2009; Cianferoni & Bartolozzi 2016; Amorim et al. 2016). This question was again raised in a letter supported by 35 signatories published in the journal Nature (Pape et al. 2016) on 15 September 2016. On 25 September 2016, the following rebuttal (strictly limited to 300 words as per the editorial rules of Nature) was submitted to Nature, which on 18 October 2016 refused to publish it. As we think this problem is a very important one for zoological taxonomy, this text is published here exactly as submitted to Nature, followed by the list of the 493 taxonomists and collection-based researchers who signed it in the short time span from 20 September to 6 October 2016

A new low altitude species of Megophrys Kuhl and van Hasselt (Amphibia: Megophryidae), from Assam, Northeast India

Mahony, Stephen, Sengupta, Saibal, Kamei, Rachunliu G. (2011): A new low altitude species of Megophrys Kuhl and van Hasselt (Amphibia: Megophryidae), from Assam, Northeast India. Zootaxa 3059: 36-46, DOI: 10.5281/zenodo.27892

Megophrys (Xenophrys) medogensis Fei, Ye and Huang 1983

Megophrys (Xenophrys) medogensis Fei, Ye and Huang, 1983 (Figure 13) Megophrys omeimontis medogensis Fei, Ye and Huang 1983:49 In: Two new subspecies of Megophrys omeimontis Liu from China (Amphibia, Pelobatidae). Acta Herpetologica Sinica, New Series, 2(2): 49–52. Holotype. Adult male (CIB 73 II0015: Figure 13), from “Motuo, Xizang, alt. 1000m ” (= Medog [or Motuo] town [~ 29°19'56"N, 95°20'27"E], Medog County, Nyingtri Prefecture, Tibet [or Xizang] Autonomous Region, China), collectors presumably Fei, Hu, Ye and Wu on 17 July 1973 (Fei et al. 1977). Paratypes [not examined]. 16 adult males, one juvenile and some tadpoles (CIB specimen numbers not provided), “collected from the same locality, alt. 850–1350m ”, collectors as above, including Huang, on 17–24 July 1973 and 21 July–5 August 1977 (Fei et al. 1977). Examined specimens. Adult male (CIB 73II 0015: images only: Figure 13), holotype. Holotype description. Refer to Fei et al. 1983 for the holotype description. For additional details of colouration and markings, see figures in Zhao et al. (2005), Fei et al. (2010, 2012) and Li et al. (2010) of specimens from the type locality. Systematic position. Refer to the Systematic position section above for M. robusta. Morphological comparison. Adult body size ranges provided for M. medogensis are from Fei et al. (2009) (adult males, N =16). Megophrys medogensis differs from M. monticola and M. zhangi by its much larger adult body size, male SVL 57.2–68.0 mm (vs. male SVL 38.2–49.5 mm, N =17; male SVL 32.5–37.2 mm, N =3, respectively); differs from M. mangshanensis by absence of white upper lip stripe (vs. present); differs from M. robusta by its smaller adult male size, SVL 57.2–68.0 mm (vs. male SVL 73.5–83.1 mm, N =6). For comparisons with additional species covered in this study, refer to relevant morphological comparison sections for those species. Etymology. The specific epithet “ medogensis ” is a toponym, derived from the type locality of Medog county. Suggested common name: Medog Horned Frog. Distribution. Megophrys medogensis is currently known with certainty from the type locality (Figure 8A) “Medog”, between 850 and 1350 m elevation (Fei et al. 1983, 2012). This species has also been reported from the southern slopes of Mt. Namjagbarwa (or Namcha Barwa) adjacent to Medog town (Zhao & Li 1987). Additional localities in Yunnan Province, China on the distribution map provided by Fei et al. (2009) are erroneous (see Remarks below). The type locality is situated near to the northern border of the Upper Siang district, Arunachal Pradesh state, Northeast India, where this species is likely to be present at similar elevation. Habitat and natural history. At the type locality, males were reported to frequent leaf litter and small streams in forests, and were heard vocalising especially on nights with clear skies (14 July–08 August). Tadpoles allocated to this species were collected from amongst rocks at the base of a waterfall of a small stream (Fei et al. 2009), but the authors did not justify how they identified the tadpoles. Remarks. In the English abstract of the original description, Fei et al. (1983) only provided a brief comparison of this species with M. omeimontis Liu, 1950 and M. jingdongensis Fei and Ye, 1983 (in Fei et al. 1983), congeners that they determined to be morphologically most similar to M. medogensis. The abstract provided the name as “ Megophrys omeimontis motuoensis ”, but elsewhere in the text the name was provided as “ Megophrys omeimontis medogensis ”, indicating that the name in the abstract was an accidental erroneous spelling (also noted by Zhao & Adler 1993). Fei et al. (1983) provided a line drawing of the profile view of the head and ventral view of the foot, but not measurements of specimens. Fei et al. (2009) provided a table of measurements for the type series, and Fei et al. (2010) provided low-resolution photos of the holotype in preservation. Zhao et al. (2005), Fei et al. (2010, 2012), and Li et al. (2010) provided images of live individuals. Fei et al. (2009) included a distribution map showing several localities in Yunnan province for Megophrys medogensis but did not provide their source (neither publications nor specimens) for these additional localities. In Fei et al. ’s (2009) book, the species Megophrys jingdongensis is accompanied by a distribution map that is identical to the one provided for M. medogensis, but without the locality “Medog” marked. Thus it is likely that the Yunnan localities for M. medogensis are the result of printing error, and should not be considered for this species’ distribution. Fei et al. (2012) provided a map for this species including only the type locality, indirectly correcting the error of Fei et al. (2009).Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 37-39, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/261020

Xenophrys gigas Jerdon 1870

Neotype for Xenophrys gigas Jerdon, 1870 and Megalophrys major Boulenger, 1908 (by present designation). Adult male (BMNH 1947.2.24.93 [rr. BMNH [18]72.4.17.399]: Figure 14), from Khasi Hills, Meghalaya state [previously “Darjeeling”––see Remarks section], Northeast India, collected by T.C. Jerdon, collection date ~1870. Examined specimens. Adult male (BMNH 1947.2.24.93), neotype; four adult males (BMNH 1947.2.24.94 [rr. BMNH [18]72.4.17.400]; BMNH 1947.2.24.95 [rr. BMNH [18]72.4.17.401]; BMNH 1947.2.24.96 [rr. BMNH [18]72.4.17.402]; BMNH 1947.2.24.98 [rr. BMNH [18]72.4.17.404]), two adult females (BMNH 1947.2.24.92 [rr. BMNH [18]72.4.17.398]; BMNH 1947.2.24.97 [rr. BMNH [18]72.4.17.403]), one unsexed subadult (BMNH 1947.2.24.99 [rr. BMNH [18]72.4.17.405]), and one juvenile (BMNH 1947.2.25.1 [rr. BMNH [18]72.4.17.406]), details as per the neotype; four adult males (ZSIC 9744; ZSIC 9756; BMNH 1908.4.8.4 [ex. ZSIC 9743]; BMNH 1908.4.8.5 [ex. ZSIC 9736]), one adult female (ZSIC 9746), and one unsexed juvenile (BMNH 1908.4.8.6 [ex. ZSIC 9760]) from “Cherrapunji [=Sohra]”, East Khasi Hills district, Meghalaya state, Northeast India, collected by J.H. Bourne, collection date unknown; two adult males (SDBDU 2007.229; SDBDU 2007.230), from the Terunhü River, Kenrunhü (25°55'4"N, 94°5'24"E, 510 m asl.), New Sendenyu Village, Tseminyu sub-division, Kohima district, Nagaland, Northeast India, collected by RGK on 0 3 October 2007; one adult female (CES 18901 [field number RGK 0073]: Figure 15A & B), from Raenghkucpaeng duithuak (duithuak = stream, in local Rongmei [Nruangmei] language), near St. Joseph School (24°51'40"N, 93°38'33"E, 470 m asl.), Khumhzi II village, Luangmai (Noney) district (erstwhile [before 2016] in Tamenglong district), Manipur state, Northeast India, collected by RGK on 23 May 2013; adult male (CES 18902 [field number RGK 0088]: Figure 15C & D), from Mpeih baek duithuak (24°51'0"N, 93°37'23"E, 450 m asl.), Khumhzi village, Luangmai district, Manipur state, Northeast India, collected by RGK on 26 May 2013; adult male (CES 18903 [field number RGK 0089]: Figure 15E & F), from near Khangchiuluanh (Khongjaron) Church (24°58'1"N, 93°29'53"E, 1255 m asl.), Khangchiuluanh village, Tamenglong district, Manipur state, Northeast India, collected by RGK on 27 May 2013. Neotype description (measurements in mm). Mature male (SVL 75.0) (Figure 14). Head moderately large, longer than wide (HW 29.0, HL 29.7, IFE 12.6, IBE 20.9); snout bluntly pointed in dorsal view, obtusely protruding in lateral view, without rostral appendage (Figure 14C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout very slightly concave; eye length ~2.5 times as long as maximum diameter of visible portion of tympanum, and shorter than snout length (EL 9.2, TYD 3.7, SL 10.6); eye–tympanum distance (TYE 6.3) longer than diameter of visible portion of tympanum; tympanum oval-shaped, obliquely orientated with upper ~20% concealed by supratympanic ridge (Figure 14C); pupil vertically elliptical; nostril positioned laterally, closer to eye than to snout tip (EN 4.2, NS 6.5); internarial distance greater than upper eyelid width, and equal to narrowest point between upper eyelids (IN 9.4, UEW 6.0, IUE 9.4); pineal ocellus not visible externally; vomerine ridges present, medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, slightly closer to choanae than to each other; vomerine teeth short; maxillary teeth present; tongue moderately large, with small notch posteriorly, medial lingual process absent. Forelimbs long, thin (Figure 14A & B), forearms moderately enlarged relative to upper forelimbs, and shorter than hand length (FAL 18.4, HAL 18.7); fingers long, narrow, without lateral fringes (Figure 14D), finger length formula IV–<” configuration, composed primarily of single row of closely spaced asperities; dorsal surfaces of forearms, thighs and shanks with short transverse ridges; posterior thighs with small scattered pustular tubercles, remaining dorsal surfaces of forelimbs and hindlimbs otherwise smooth; gular region, chest, abdomen and ventral surfaces of limbs smooth; pectoral glands small, raised, level with axilla on chest; femoral glands large, flat, on posterior surface of thighs, slightly closer to knee than to cloaca; distinct pair of enlarged glands present adjacent to cloaca, one on each side, that are smaller than femoral glands but larger than surrounding tubercles; small black and white dermal asperities present, forming broad dense band circummarginally on gular region, dense on upper lips, lower loreal region, and tympanic region (especially dense on tympanum), moderately dense on dorsal surface of eyelids, entire dorsal surface of head and back, increasing in density posteriorly and on dorsal ridges, absent from all remaining surfaces. Colouration: In preservative (Figure 14): Dorsal and lateral surfaces of body, and dorsal and lateral surfaces of head primarily brown; light-edged, faintly darker brown triangular marking between eyes; no X-, Y-, or V-shaped markings on dorsum; tubercles primarily white on posterior flanks only; lateral surfaces of head below supratympanic ridges and canthus rostralis dark brown with cream-white stripe on upper lip, extending from nostril to rear of jaw, bordered below by thin dark brown stripe; outer edges of eyelids dark brown; dorsal and lateral surfaces of forelimbs and hindlimbs primarily mid brown; three dark brown blotches on dorsolateral surface of forearms; dorsal surface of outer three fingers with faint brown blotches; dorsal surfaces of hindlimbs without distinct transverse crossbars; lateral surfaces of thighs and shanks with dark brown spots and blotches; throat and chest primarily plain mid brown with few cream-white spots along edge of gular region, light-edged wide dark brown stripe extending from posterior edge of mandible onto base of forearms; abdomen, ventral surfaces of forelimbs, thighs, shanks, and dorsal surfaces of tarsi and feet immaculate pale brown; some small dark brown spots and blotches laterally along lower flanks on both sides bordering lower-most tubercles; area surrounding vent and posterior surfaces of thighs primarily mid to dark brown with small cream-white tubercles; ventral surfaces of tarsi and feet pale greyish-brown; hands ventrally pale greyish-brown; pectoral and femoral glands creamish-white. In life: Not documented for neotype. Refer to Figure 15 for examples of colour in life of referred specimens. Variation. Refer to Table 1 for morphometric variation within the referred specimen series, consisting of 12 adult males, three adult females, two subadults, and a juvenile. Referred specimens generally resemble neotype for most morphological characters with the following exceptions: Webbing between digits varies slightly between individuals and sexes, i.e., webbing formula on Toe IV: males from 3.2IV3.2 to 3.6IV3.6, females 4IV 4.3; lateral fringes on toes vary from moderately weak from bases to tips on all toes, to almost completely absent on some individuals; relative finger lengths seem to vary between populations, IV<II=I<III on all “Darjeeling” specimens except a subadult (BMNH 1947.2.24.99) with IV=II=I<III, all Sohra specimens have IV=II=I<III, except a female (ZSIC 9746) with IV<II<I<III, and all aforementioned variations, with an additional IV<I<II<III for RGK 0 0 98, are observed in the Nagaland-Manipur specimens; vomerine ridges vary from sub-circular to ovoid, positioned slightly closer to choanae than to each other on many individuals; posterior edge of tongue on some specimens appears more deeply notched than others, likely due to preservation condition; dorsolateral ridges typically weak to moderately well developed, varying in length from ~60 to 90% trunk length; between ~5 and 30% of tympanum can be concealed by supratympanic ridge; dermal tubercle cover on flanks varies considerably, some with only sparse scattering of small tubercles (e.g., Figure 15A), others with moderately dense cover of heterogeneous sized (large to small) tubercles (e.g., Figure 15C & E); dorsal markings vary from faint to distinct complete X-shaped, Yshaped, hourglass-shaped marking, or just a V-shaped marking on parietoscapular region, not visible on some older specimens presumably due to fading; some specimens have faint spotting and blotching on chest and abdomen (e.g., Figure 15F); density and surface coverage of dermal asperities varies extensively among male specimens, if present on females, asperities sparse on mid-dorsum increasing in density posteriorly to above cloaca, but absent from other surfaces, asperities typically absent on juvenile and subadult specimens examined; white upper lip stripe not present on metamorphs and juveniles (with SVL <50 mm), instead lateral surfaces of head plain dark brown. Secondary sexual characters. Males: nuptial pads present, weakly raised, covered with brown/black microasperities on freshly collected specimens (appear as microgranules on old specimens), primarily covering dorsal surface of base of Finger I, narrowing distally, extending to mid-proximal phalange on inner dorsal side; nuptial pad on Finger II small, oval, positioned on base of digit on inner dorsal side (extending to mid-proximal phalange on some individuals); external vocal sac indistinct; internal vocal slit present on floor of mouth near rear of mandible, one on each side; forearms enlarged relative to upper forelimbs. Females: mature ova without pigment; nuptial pads, vocal sac, vocal slits, and enlarged forearms, all absent. Morphological comparison. Megophrys major s.s. (adult males, N =12, adult females, N =3) differs from M. monticola and M. zhangi by its larger adult body size, male SVL 71.6–87.5 mm, female SVL 85.6–98.2 mm (vs. male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6; male SVL 32.5–37.2 mm, N =3, respectively); differs from M. robusta and M. medogensis by presence of distinct continuous whitish-cream upper lip stripe (vs. absent), by basal webbing present (vs. absent); differs from M. mangshanensis by basal webbing between toes present (vs. absent), lateral fringes on toes occasionally present (vs. absent), larger adult body size, male SVL 71.6–87.4 mm, female SVL 85.6–98.2 mm (vs. male SVL 62.5 mm, N=1, female SVL 73 mm, N=1). For comparisons with subsequent species covered in this study, refer to relevant morphological comparison sections for those species. Systematic position. OTU 5 (“ M. cf. major 3”) in the molecular analyses of this study and Mahony et al. (2017) represents Megophrys major s.s. The systematic position of M. major s.s. within the MMC clade is not fully resolved (Figures 2–5; Appendix I, Table 3; Appendix II, Figures 1, 2 & 5: as M. cf. major 3/OTU 5). It was consistently found to be the sister taxon to a new species (OTU 7/“ M. cf. major 1”) described below, but it is not yet clear which species these sister taxa are most closely related to within the MMC. Uncorrected p -distance for the 16S rRNA gene between these sister taxa was 6.4–6.5% (Appendix I, Table 6). Etymology. The original species epithet “ gigas ” is derived from the Greek word meaning “giant”, later replaced with “ major ”, Latin for “large”. Suggested common name: Jerdon’s White-lipped Horned Frog. Distribution. This species was considered widespread from Northeast India to southeastern Vietnam but M. major sensu lato (s.l.) is now known to represent a species complex (Chen et al. 2017; Mahony et al. 2017; this study). Only the following localities (between 450 and 1255 m asl.) represent the verified distribution of M. major s.s. (Figure 8A): Sohra [=Cherrapunji], in the East Khasi Hills district, Meghalaya state; Khumhzi, in Luangmai district and Khangchiuluanh, in Tamenglong district, Manipur state; and Tseminyu, in Kohima district, Nagaland. The locality “Darjeeling” associated with Jerdon’s BMNH series of specimens is considered erroneous (see Remarks for details). The eastern distribution limit for M. major s.s. is currently unclear, but it might be expected to extend into adjacent areas of western Myanmar. A report of this species from Tura Peak, West Garo Hills district in Meghalaya state was provided by Sangma and Saikia (2015). The authors provided no diagnostic characters for their specimens so the taxonomic identity of animals from this locality warrants careful investigation considering our study identifies the presence of three MMC species in Meghalaya state. Habitat and natural history. There have been many observations and notes published regarding M. major s.l., primarily from Southeast Asian countries (e.g., Bain & Nguyen 2004; Ohler et al. 2000; Stuart 1999, 2005), which are likely to represent different species. Some old reports of M. major tadpoles (e.g., Annandale 1912) from Northeast India are not considered reliable, since most tadpoles were identified by association of adult frogs present in the collection stream, not accounting for the fact that in our experience most streams in Northeast India are inhabited by two or more Megophrys species. Based on our observations of recently collected specimens, a female (CES 18901) collected towards the end of May contained mature ova, and males were found calling during May (CES 18902, CES 18903) and October (SDBDU 2007.229, SDBDU 2007.230), indicating that the breeding season for this species probably extends from the pre-monsoon (May) until post monsoon (October). SDBDU 2007.229 and SDBDU 2007.230 were found calling amongst fallen and dead reeds on the banks of the Terunhü River in Nagaland at around 18:30 h (just after dusk). The Terunhü River at the collection locality is a moderately fast flowing stream passing through a disturbed secondary forest patch. CES 18902 was found calling during the daytime perched ~ 2.3 m above ground level on a leaf of a wild cardamom on the bank of the Mpeih bak duithuak (stream). CES 18903 was found calling from a hole in the retaining wall where a clear water stream exits an underground diversion. The gravid female (CES 18901) was collected nearby an unnamed stream. No further individuals could be located along that stream during the subsequent two nights of searching, indicating that breeding activity had not yet reached a peak (or was experiencing a lull). Remarks. On Xenophrys gigas Jerdon, 1870: The original type series of Xenophrys gigas consisted of five syntype specimens in Jerdon’s personal collection collected from “ Sikim [sic] and the Khasi Hills” (Jerdon 1870). Soon after the description of Jerdon’s X. gigas, Anderson (1871b) stated “I have a specimen of Dr. Jerdon’s X. gigas beside me [in the Indian Museum (now ZSIC)], but I find that in no way differs from the Darjeeling large specimens, which are the adults of this species [Xenophrys monticola Günther, 1864]”. With that statement, X. gigas was regarded to represent a junior synonym of X. monticola for the subsequent four decades. Sclater (1892b:33) listed all Indian Museum amphibian specimens by numbers, localities and collectors, which included seven specimens of “ X. monticola ” sensu Anderson (1871b) deposited by Jerdon, six from “Darjeeling” (ZSIC 9673–9677, ZSIC 9721) and one from “Khasia Hills” (ZSIC 9679). We know now (discussed below) that the large species from Darjeeling that Anderson (1871b) was referring Jerdon’s X. gigas specimen to is M. robusta. However, it is not clear whether that particular specimen was one of Jerdon’s five syntype specimens of X. gigas. Deuti et al. (2017) referred five of the aforementioned “Darjeeling” specimens (ZSIC 9673–9677), and three others not mentioned by Sclater (1892b; ZSIC 9722–9724) to Megophrys sanu comb. nov. (as juveniles). The taxonomic identities of ZSIC 9721 and ZSIC 9679 require clarification. The whereabouts of the original type series of Xenophrys gigas, and which specimens actually belong to the original type series of this species, is an imbroglio. Chanda et al. (2000) referred four specimens (ZSIC 9670, ZSIC 9681, ZSIC 10777, ZSIC 10779) as types of X. gigas Jerdon, from “Darjeeling”, West Bengal, India. ZSIC also have an additional two specimens from Darjeeling that they regard to be the types of X. gigas (ZSIC 9650 and ZSIC 9668) (SM per. obs. during a visit to the ZSIC in 2010). Several problems exist with the recognition of these six ZSIC specimens as the types for this species: 1) all of these specimens were collected by J. Gammie (Sclater 1892b:33 [as Leptobrachium monticola]); Jerdon (1870) clearly stated that he “obtained” the type specimens from the type localities—Jerdon acknowledged throughout this paper when specimens were sent to him by other collectors, so it may be interpreted that he (or his party) were the collectors of the specimens; 2) ZSIC 9650 (SVL 51.1 mm) is a gravid adult female M. monticola (SM per. obs.). ZSIC 9668 was not examined in detail but is either an adult M. monticola or a juvenile M. robusta ––both specimens do not fit Jerdon’s original description by being “large”, presumably relative to the considerably smaller Xenophrys monticola, the only other Xenophrys species known at that time; 3) ZSIC 10777 and ZSIC 9681 are M. robusta (SM per. obs.); 4) Sclater (1892a, 1892b) provided a list of types in the Indian Museum that included only one megophryid, Scutiger (as Cophophryne) sikkimensis (Blyth, 1855), and Sclater (1892a) explicitly stated that the collections then contained the types of only one of Jerdon’s species, Hyla annectans (Jerdon, 1870). Before leaving India, Jerdon visited and collected specimens in and around Darjeeling–Sikkim and the Khasi Hills, but he notoriously did not maintain notes on his collections, relying largely on memory for details of localities (Günther 1875). Upon his return to England, Jerdon assisted Günther with the accessioning of his collection into the BMNH– –“The work of systematically arranging and naming this collection was carried on jointly by him [Jerdon] and myself, and proceeded as far as the genus Tropidonotus, when it was interrupted by an illness from which he never recovered….and I am therefore ignorant of the habitat [collection localities] of a part of the specimens which were still unexamined at the time of his death” (Günther 1875). Jerdon’s original “ X. gigas ” collection consisted of a series of presumably 17 specimens (BMNH [18]72.4.17.398–414), and was one of the last taxa from his collection to be accessioned (BMNH Accession Register). The BMNH Accession Register gives the locality “Darjeeling (& Khasia)”, but the locality appeared in subsequent literature as Darjeeling only (e.g., Boulenger 1882 [“h”]). Doubts regarding whether some of Jerdon’s other herpetological specimens were from the Khasi Hills or from Darjeeling have been noted elsewhere (e.g., the type locality of Ixalus jerdoni Günther, 1876 [= Frankixalus jerdoni] from “Darjeeling” was subsequently found to be widespread in the Khasi Hills [Biju et al. 2016] but not reported from Darjeeling or surrounding regions since its original description). Jerdon deposited most (if not all) of his type specimens in the British Museum and Indian Museum, but since the Indian Museum did not (according to Sclater [1891a, 1891b]) contain Jerdon’s megophryid type specimens, the BMNH specimen series may have contained some, or all, of the five original specimens based on which Jerdon (1870) coined the name. Remarks on Megophrys major Boulenger, 1908: Xenophrys gigas was subsequently transferred to the genus Megalophrys by Boulenger (1908) rendering Megalophrys gigas (Jerdon, 1870) a junior homonym of Megalophrys gigas Blyth, 1855 [now Nanorana liebigii (Günther, 1860)]. To correct this, Boulenger (1908) coined the replacement name Megalophrys major, and provided a morphological description of the species with a note that Jerdon (1870) named X. gigas based on specimens from Darjeeling and the Khasi Hills “now preserved in the British Museum [now NHMUK] and Indian Museum [now ZSIC]”. Considering Sclater (1892a, 1892b) did not recognise types for this

Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species

Mahony, Stephen, Kamei, Rachunliu G., Teeling, Emma C. (2018): Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species. Zootaxa 4523 (1): 1-96, DOI: https://doi.org/10.11646/zootaxa.4523.1.

Ichthyophis moustakius Kamei, Wilkinson & Gower, 2009, sp. nov.

&lt;i&gt;Ichthyophis moustakius&lt;/i&gt; sp. nov. &lt;p&gt;(Figs. 1, 4&ndash;5; Tables 1&ndash;2)&lt;/p&gt; &lt;p&gt; &lt;b&gt;Holotype&lt;/b&gt;. BNHS 5213, an adult female, collected by SDB and RGK on 3 June 2006 from Aziuram duikhun (duikhun = a pond) (25&deg;01&rsquo;43&rsquo;&rsquo;N, 93&deg;24&rsquo;51&rsquo;&rsquo;E; 990 m asl), Aziuram village, Tamenglong district, Manipur, India.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Paratypes&lt;/b&gt; (n = 4). BNHS 5214&ndash;5216, three adult females, and BNHS 5219, an adult male, collected along with holotype.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Referred material&lt;/b&gt; (n = 3). BNHS 5218, an adult female collected on 30 May 2008 from Guigailuang, Nriangluang namdaih (namdaih = a large village), (24&deg;59&rsquo;32&rsquo;&rsquo;N, 93&deg;29&rsquo;38&rsquo;&rsquo;E; 1107 m asl), Tamenglong district; SDB 1513, an adult female collected on 7 June 2007 from Nswanram village (25&deg;01&rsquo;04&rsquo;&rsquo;N, 93&deg;34&rsquo;09&rsquo;&rsquo;E; 838 m asl), Tamenglong district; BNHS 5220, an adult male collected on 29 May 2008 from Duidip Chaengluan (24&deg;55&rsquo;54&rsquo;&rsquo;N, 93&deg;24&rsquo;53&rsquo;&rsquo;E; 306 m asl), Bamgaizaeng village, Tamenglong district. Specimens were collected by SDB and RGK.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Diagnosis&lt;/b&gt;. &lt;i&gt;Ichthyophis&lt;/i&gt; with broad (W/S &lt;4) fairly regular, mostly solid, lateral yellow stripes from anterior of tail to at least CM, broad along mandibles with narrow anterior gap, expanded and visible ventrally on collars, connected by spurs to disc; arched yellow stripes extending between nares and TAs, broader at former than latter; not known to attain lengths greater than 300 mm, 18 &lt;L/W &lt;25; fewer than 300 AGs, paler than adjacent skin; head somewhat more U-than V-shaped, fairly short (25&gt; L/H&gt; 19); TAs about twice as far from nares than from eyes, variable (1.9 &lt;TN/TE &lt;2.3); collars of similar lengths; scales absent on collars, beginning from about fourth or fifth annulus, with five rows posteriorly on dorsum; similar numbers of IMs and dentary teeth.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Description of holotype&lt;/b&gt;. Some morphometric and meristic data given in Table 1. Good condition, several annular scale pockets opened, c. 15 mm midventral longitudinal incision c. 65 mm anterior to vent. 105 vertebrae. Head, nuchal region and trunk slightly dorsoventrally compressed. Girth maximal near midbody, decreasing gradually, more substantially over posteriormost c. 10 mm, L/W = 20.6. Tail upturned towards tip. Head short (L/H = 21.7). In lateral view, CM equidistant from top and bottom of head. In ventral view, lower jaws inset from upper, slightly more so anteriorly than at level of TAs. Eye closer to top of head than to lip in lateral view, surrounded by narrow whitish ring, eye diameter c. 0.6 mm, lens no larger than naris. TAs slightly smaller than eyes, larger than nares, less than twice as distant from nares than from eyes, very close to lips, just below (touching) imaginary lines between nares and CMs. Nares slightly posterior to level of anterior margin of mouth; in lateral view equidistant from tip, top and bottom of snout, inset a little less than one naris diameter from sides in dorsal view. Teeth very slender, strongly recurved, dentary series extend about one third the length of the IM series (with 5 teeth on each side) behind posteriormost IM. Tongue not strongly plicate, margin overlying all except anteriormost IMs. Choanae narrow, distance between them four or five times their greatest width. Collar region slightly more massive than adjacent head and body, delimited by weak constrictions, lacking TGs. C2 slightly longer than C1 measured laterally. Dorsally, NG3 complete with gentle anteromedial curvature.&lt;/p&gt; &lt;p&gt;AGs mostly ventrally incomplete on anterior one third, four interrupted in disc region, four complete ventrally on tail, last two incomplete dorsally. Dorsally, anteriormost c. 25 AGs and posteriormost nine curve slightly anteromedially. Ventrally, anterior AGs strongly angulate, degree of posteromedial curvature about length of two annuli up to midbody, decreasing posteriorly, more or less orthoplicate by 27th annulus anterior to disc. Tiny scales scattered in first AG, three rows of small (0.9 x 0.5 mm) scales in shallow pockets as far anterior as the 11th annulus. At midbody four rows of larger (up to c. 1.4 x 1.0 mm) scales in deep pockets dorsally, three or four rows of smaller scales (up to c. 1.1 x 0.5 mm) in shallow pockets ventrally. Posteriorly (c. 20 annuli anterior to vent) five rows of more rounded scales (up to c. 1.1 x 1.0 mm) in deep pockets dorsally, smaller scales (1.0 x 0.8 mm) in four rows in shallow pockets on venter. No papillae on disc. Fifteen irregular denticulations around vent.&lt;/p&gt; &lt;p&gt;In preservation, lilac-grey with brownish tinges, slightly paler ventrally. Lateral stripes extending from about second or third posteriormost annulus to about eye level on upper jaws and close to mandible tips, expanding ventrally on collars, especially C1, narrowly broken just behind NG1. Narrow yellow lines extending dorsally then curving anteriorly from TAs to nares, thickest near nares; AGs paler, especially posteriorly. In life, dorsum uniformly dark reddish grey, venter pale reddish grey; lateral stripe bright yellow, vent disc mauvish.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Variation&lt;/b&gt;. Morphometric and meristic data for the type series and referred specimens are given in Table 1, and summarised in Table 2. There are between three and six dentary teeth posterior to the last IM on each side. The distance between the choanae ranges from three to five times the maximal choanal width. Ranges of the numbers of AGs anterior to the vent that are orthoplicate, and of anterior and posterior AGs that curve anteromedially on the dorsum are quite small (21&ndash;29, 8&ndash;26, and 7&ndash;13, respectively). The number of denticulations around the vent ranges from 13 to 17. Compared to the holotype, the paratypes have mostly &lt;i&gt;Ichthyophis khumhzi Ichthyophis moustakius Ichthyophis sendenyu&lt;/i&gt; continued next page incomplete AGs ventrally on a larger proportion of the body. Stripes may or may not be broken across the collars, and may extend as far as the TAs. Papillae on the disc are present in one of the two male specimens (BNHS 5220). The tail is upturned in only a single paratype (BNHS 5215).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Etymology&lt;/b&gt;. The species name is derived from the Greek word &lt;i&gt;moustakius&lt;/i&gt;, meaning moustache, referring to the distinctive yellow, arched stripes extending between the TAs and nares. The specific epithet is considered to be a noun in apposition.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Suggested common name&lt;/b&gt;. Manipur moustached Ichthyophis.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Distribution and natural history&lt;/b&gt;. Aziuram, Nswanram, Nriangluang, and Bamgaizaeng in Tamenglong district, Manipur (Fig. 1), at c. 300 to 1100 m asl. All specimens from the type series were collected by digging in perennially wet soil in the heart of the village. Four females with egg clutches containing 9&ndash; 19 eggs per clutch in early developmental stages were observed in the dry period (first week of June), suggesting that this species is a relatively early breeder compared with many/most of the frogs in the same region (SDB pers. obs.). BNHS 5220 was collected from underneath a rock in a secondary forest.&lt;/p&gt;Published as part of &lt;i&gt;Kamei, Rachunliu G., Wilkinson, Mark &amp; Gower, David J., 2009, Three new species of striped Ichthyophis (Amphibia: Gymnophiona: Ichthyophiidae) from the northeast Indian states of Manipur and Nagaland, pp. 26-42 in Zootaxa 2267&lt;/i&gt; on pages 32-37, DOI: &lt;a href="http://zenodo.org/record/190862"&gt;10.5281/zenodo.190862&lt;/a&gt

Megophrys (Xenophrys) major Mahony & Kamei & Teeling 2018, s.s.

Megophrys (Xenophrys) major s.s. Boulenger, 1908 (Figures 14 & 15; Table 1) Xenophrys gigas Jerdon 1870:85 (partim: Khasi Hills). In: Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, March, 1870: 66–85. Megalophrys major Boulenger 1908:410, 416, Pl. xxiii. In: A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908: 407–430 + Pl. xxii–xxv + fig. 71.Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on page 39, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/261020

Megophrys (Xenophrys) periosa Mahony & Kamei & Teeling 2018, sp. nov.

Megophrys (Xenophrys) periosa sp. nov. (Figures 22 & 23; Table 1) Holotype. Adult male (BNHS 6055 [field no. SDBDU 2009.793]: Figures 22, & 23A & H), from Pangin town (28°12'33.96"N, 94°59'10.02"E, 450 m asl.), East Siang district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 26 July 2009. Paratypes. Four adult males (BNHS 6057 [field no. SDBDU 2009.794]: Figure 23B; BNHS 6058–6060 [field nos. SDBDU 2009.1170 – 1172]: Figure 23H), collected along with the holotype by members of the Systematics Lab, University of Delhi on 26–28 July 2009; four adult males (BNHS 6061 [field no. SDBDU 2009.1243]: Figure 23E; BNHS 6062 [field no. SDBDU 2009.1244], BNHS 6063–6064 [field nos. SDBDU 2009.1265 – 1266]: Figure 23G), and one adult female (BNHS 6056 [field no. SDBDU 2009.1285]: Figure 23F), from the Sessa River, nearby Sessa village (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009. Referred specimens. Two adult males (SDBDU 2009.1189: Figure 23D; SDBDU 2009.1190: Figure 23C), from Rigo Village (28°9'34.56"N, 94°47'19.38"E, 260 m asl.), Along town, West Siang district, Arunachal Pradesh state, Northeast India, collected by Systematics Lab members on 30 July 2009; one adult male (SDBDU 2009.1267), from the Sessa River, nearby Sessa village (27°6'4.02"N, 92°31'38.52"E, 1110 m asl.), West Kameng district, Arunachal Pradesh state, Northeast India, collected by members of the Systematics Lab, University of Delhi on 07–09 August 2009. Provisionally referred specimens (see Remarks). One juvenile male (SDBDU 2009.132), and two juvenile females (SDBDU 2009.133 & SDBDU 2009.134), from Pangkava ju (24°40'21.6"N, 94°28'19.86"E, 820 m asl.), Kangpat Khullen, Kamjong sub-division, Ukhrul district, Manipur state, Northeast India, collected by RGK and SDB on 21 May 2009; adult female (CAS 232938 ––tissue only), from Hepu Stream (25°5'25.2"N, 96°24'13.2"E), Hepu village, Moe Nyin township, Myitkyina district, Kachin state, northern Myanmar, collected by G.O.U. Wogan, J.A. Wilkinson, J.V. Vindum, H. Win, T. Thin [“Additional collectors: K.S. Lwin, A.K. Shein and H. Tun ”] on 14 May 2003. Holotype description (measurements in mm). Mature male (SVL 93.8) (Figures 22, & 23A & H). Head moderately large, wider than long (HW 35.0, HL 33.6, IFE 15.5, IBE 25.3); snout bluntly pointed in dorsal view, obtusely protruding beyond mandible in lateral view, without rostral appendage (Figure 22C); loreal region acute, concave; canthus rostralis angular; dorsal surface of snout very slightly concave; eye diameter twice maximum diameter of visible portion of tympanum, and shorter than snout length (EL 10.7, TYD 5.4, SL 11.7); eyetympanum distance (TYE 8.2) less than twice diameter of visible portion of tympanum; tympanum oval-shaped, oblique (Figure 22C), with upper ~15% concealed by supratympanic ridge; pupil vertically elliptical; nostril positioned laterally, closer to eye than to snout (EN 5.2, NS 7.1); internarial distance greater than eyelid width, and equal to narrowest point between upper eyelids (IN 10.5, UEW 8.7, IUE 10.5); pineal ocellus not visible externally; vomerine ridge present, medium sized, ovoid, moderately raised, orientated acutely, positioned between to slightly posterior to choanae, equidistant from choanae and each other; vomerine teeth short; maxillary teeth present; tongue not observed due to fixation of jaw and in interest of preventing potential damage to jaws by forcing open mouth wide enough for examination. Forelimbs long, thin (Figures 22A & B, & 23A), forearm moderately enlarged relative to upper forelimb, and shorter than hand length (FAL 20.6––left side; right side deformed; HAL 24.0); fingers long, narrow, without lateral fringes (Figure 22D), finger length formula IV –75% of the trunk length (Figure 23 A–F); typically only upper border of tympanum is concealed by supratympanic ridge, though up to ~25% of tympanum appears to be concealed on some specimens; parietoscapular-sacral ridge configuration varies considerably amongst individuals, i.e., “>- (”, “>–– |”, “>- <”, or only V-shaped parietoscapular ridge present; coverage of dermal asperities varies mostly in density between individual males in comparison with holotype, however, on some specimens both black (or brown) and white asperities were observed, may sparsely cover upper lips, loreal region, front of snout, posterior dorsal surface of upper eyelids, head, and dorsal surface of tibia-tarsus joints; asperities white on female, restricted to dorsal surface of body, sparse on middorsum, increasing in density posteriorly to above cloaca; outer metacarpal tubercles not visible on most specimens; tubercle cover on flanks varies considerably, some with only sparse scattering of small tubercles, others with moderately dense cover of heterogeneous (large to small) sized tubercles (Figure 23 A–F). Dorsal and ventral markings vary considerably between individuals (see Figure 23 which represent extremes in variation). Secondary sexual characters. Males: nuptial pads present, weakly raised, covered with brown/black microasperities on freshly collected specimens, covering most of dorsal surface of Finger I; nuptial pad on Finger II medium sized, oval, positioned on base of digit on inner dorsal side, extending almost to base of distal phalange; external vocal sac indistinct; large internal vocal slits present on floor of mouth near rear of mandible, on each side; forearms enlarged relative to upper forelimbs. Female: mature ova without pigmented poles; nuptial pads, vocal sac, vocal slits, enlarged forearms, all absent. Morphological comparison. Characters used for comparing Megophrys periosa sp. nov. with its congeners do not include those from the provisionally assigned referred specimens. Megophrys periosa sp. nov. (adult males, N =12, adult female, N =1) differs from M. medogensis, M. zhangi and M. monticola by its larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 57.2–68.0 mm, N =17; male SVL 32.5–37.2 mm, N =3; male SVL 38.2–49.5 mm, N =17, female SVL 40.5–56.1 mm, N =6, respectively); differs from M. robusta by absence of black dermal asperities on posterior abdomen of adult males (vs. usually present), Finger II<I in length, N =13 (vs. Finger I=II, N =10); differs from Megophrys flavipunctata sp. nov., Megophrys oreocrypta sp. nov., M. major s.s. and M. mangshanensis by absence of distinct white, cream or light coloured stripe along upper lips (vs. present); further differs from Megophrys flavipunctata sp. nov. and M. mangshanensis by its larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 56.9–68.4 mm, N =4, female SVL 68.0– 74.6 mm, N =3; male SVL 62.5 mm, N =1, female SVL 73 mm, N =1, respectively); further from M. major s.s. by dark spots associated with flank tubercles typically present (vs. absent), toe tips not expanded relative to adjacent toe width (vs. expanded); differs from Megophrys himalayana sp. nov. by its typically larger adult body size, male SVL 71.3–93.8 mm, female SVL 112.0 mm (vs. male SVL 68.0– 73.5 mm, N =7, female SVL 83.9 mm, N =1), typically smaller eye diameter to snout length ratio for males, ED/ES 76.6–91.5%, mean 84.3 ± 4.2% (vs. ED/ES 87.6–101.1%, N =7, mean 94.2 ± 4.6%). Systematic position. This taxon represents M. cf. major 4 (OTU 12) in the molecular analyses, and “ M. cf. major [5]” in Mahony et al. (2017). Megophrys periosa sp. nov. consistently formed a sister taxa relationship with two populations found east of the Brahmaputra River in Manipur and Myanmar referred to herein as Megophrys cf. periosa (M. cf. major 6/OTU 14 in molecular analyses). Uncorrected p -distances for the 16S rRNA gene between Megophrys periosa sp. nov. and M. cf. periosa was 1.6–2.0% (Appendix I, Table 6). The systematic position of this clade within the MMC was not fully resolved (see Systematic position section for Megophrys himalayana sp. nov. for further details), however most analyses placed this species as the sister taxon of Megophrys himalayana sp. nov. (Figures 2 & 4; Appendix I, Table 3; Appendix II, Figures 1, 2 & 5; Mahony et al. 2017). Refer to Appendix I, Table 6 for uncorrected p - distances for the 16S rRNA gene between Megophrys periosa sp. nov. and other MMSG species. Etymology. The species epithet “ periosa ” is a Latinised Greek adjective meaning “immense”, in reference to the very large size that this species attains, even compared to otherwise generally large species in the MMC. Suggested common name: Giant Himalayan Horned Frog. Distribution. Megophrys periosa sp. nov. is so far confirmed from three localities in the state of Arunachal Pradesh, Northeast India. The currently known distribution ranges from East Siang district, west through West Siang district, to at least West Kameng district, between 260 and 1110 m asl. (Figure 8A). An improved and more extensive sampling is necessary to define the east-west limits of this species’ geographic range. Specimens assigned here to Megophrys cf. periosa were collected from Ukhrul district (at 820 m asl.), Manipur state, Northeast India, and from Myitkyina district, Kachin state, northern Myanmar (Figure 8A). Habitat and natural history. All males were in breeding condition, and the female contained well-developed ova within her ovaries, indicative that the breeding season extends at least through late-July and early-August. All specimens were collected after dusk, typically perched on the rocky banks of small to large (1–15 m wide) moderately fast flowing mountain streams flowing through dense mature secondary/primary forest (e.g., Figure 9A & F). Males spaced themselves along the banks of the streams at least 10 m apart. Vocalisations typically consisted of a succession of a few notes followed by an extended silence, but were rarely heard. Healed injuries were observed on two specimens; the right hand of the holotype appears to have been badly dislocated or broken (Figure 22B & E). This large mature male had noticeably less well-developed nuptial pads than the other males, possibly as a result of lower overall fitness, although it otherwise appeared to be in good health. One other specimen (BNHS 6064) had a large swelling on Finger II on the right hand, the cause for which is unclear. Remarks. Specimens reported as M. lateralis and/or M. robusta by Borah and Bordoloi (2001), Bordoloi et al. (2000) and Sarkar and Ray (2006) from Arunachal Pradesh may apply to this species, and/or Megophrys himalayana sp. nov. The specimens reported in these studies should be re-examined to determine the correct identities. Populations provisionally assigned to this taxa (as M. cf. periosa) from eastern Manipur state, Northeast India and northern Myanmar, were found to be the sister taxon to Megophrys periosa sp. nov. (Mahony et al. 2017 [as M. cf. major [6]]; Figures 2, 4 & 5). Regardless of BPP results indicating that this lineage may represent a distinct species-level taxon (Appendix I, Table 5), we recommend that comparable series of adult specimens be studied to further determine the taxonomic status of this lineage. For now, it may provisionally be regarded as conspecific with Megophrys periosa sp. nov.Published as part of Mahony, Stephen, Kamei, Rachunliu G. & Teeling, Emma C., 2018, Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species, pp. 1-96 in Zootaxa 4523 (1) on pages 59-63, DOI: 10.11646/zootaxa.4523.1.1, http://zenodo.org/record/261020

FIGURE 11 in Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species

FIGURE 11. Megophrys robusta lectotype: adult female (BMNH 1947.2.25.19: SVL 102.0 mm) in preservation: A. dorsal view; B. ventral view; C. profile view of head; D. ventral view of foot; E. ventral view of hand

FIGURE 22 in Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species

FIGURE 22. Megophrys periosa sp. nov. holotype: adult male (BNHS 6055: SVL 93.8 mm) in preservation: A. dorsal view; B. ventral view; C. profile view of head; D. ventral view of hand; E. ventral view of foot