identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
05278791FFC6FFACEEDB04982D56FDCD.text	05278791FFC6FFACEEDB04982D56FDCD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scutiger khumbu Hofmann & Ohler & Baniya & Dubois & Flecks & Jablonski & Schmidt & Dufresnes 2025	<div><p>Scutiger khumbu sp. nov.</p><p>urn:lsid:zoobank.org:act: 3F5B9602-4B6E-467E-9AF3-77D66724ACB7</p><p>Figs 6–10; Table 3; Supp. file 1J, L–O</p><p>Diagnosis</p><p>Scutiger khumbu sp. nov. is assigned to the genus  Scutiger based on morphological characteristics and its phylogenetic position (Fig. 2). It can be distinguished based on multiple nucleotide substitutions in mitochondrial COI, cytb, and 16S sequences, as well as private alleles at the nuclear genes bfib7, ccnb2 and rag1. It can be distinguished from all other congeners by the following combination of morphological features: medium body size (male 41.6–51.1 mm SVL, female 49.4–55.6 mm; Table 3, Supp. file 1J); head wider than long; vomerine and maxillary teeth absent; vocal sac absent; relative length of fingers 3&gt; 4&gt; 2&gt; 1; small black nuptial spines on dorsal surface of first and second fingers, and inner side of third finger in males in breeding condition; inner forearm without spines; relative length of toes 4&gt; 3&gt; 5 ≥ 2 ≥ 1; subarticular tubercles absent; toes unwebbed or rudimentary webbed; a pair of pectoral glands and a pair of axillary glands with small black spines on both glands; abdomen/belly without spines; tubercles on dorsal and lateral surfaces of body (often arranged in lines) and upper limbs; tubercles conical, each tubercle with keratinized tips or 1–2 black spines in breeding condition, entire cone with horned, black coating, studded with small black tips, shimmering like black mica schist when moistened in the preserved state, in particular in specimens from the Khumbu region (Fig. 6). A triangular or Y-shaped dark brownish pattern can be present on dorsal side of the head (Supp. file 1L–M).</p><p>Etymology</p><p>The specific epithet  ʻ khumbu ʼ is a noun in apposition referring to the Khumbu Himal (also called the Everest region) in Nepal and thus remains unchanged regardless of the gender of the genus.</p><p>Type material</p><p>Holotype</p><p>NEPAL • ♂; Bagmati Province, Dolakha District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.275&amp;materialsCitation.latitude=27.696667" title="Search Plazi for locations around (long 86.275/lat 27.696667)">Jiri</a>; 27°41′48″ N, 86°16′30″ E (WGS 84); 3396 m a.s.l.; Jun. 1997; local resident leg.; ZFMK 104174.</p><p>Paratypes</p><p>NEPAL • 1 subadult; Bagmati Province, Ramechhap District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.340004&amp;materialsCitation.latitude=27.595278" title="Search Plazi for locations around (long 86.340004/lat 27.595278)">Gokulganga</a>; 27°35′43″ N, 86°20′24″ E (WGS 84); Jun. 1997; local resident leg.; ZFMK 104175  •   13 ♂♂; Solu Khumbu District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.80222&amp;materialsCitation.latitude=27.575834" title="Search Plazi for locations around (long 86.80222/lat 27.575834)">Najing Dingma</a>; 27°34′33″ N, 86°48′08″ E (WGS 84); 1 Jul. 1973; Alain Dubois and Dominique Payen leg.; MNHN 1977.1246 to MNHN 1977.1250, MNHN 1977.1252, MNHN 1977.1254, MNHN 1977.1256 to MNHN 1977.1261  •  4 ♀♀; same data as for preceding; MNHN 1977.1251, MNHN 1977.1253, MNHN 1977.1255, MNHN 1977.1262 •   3 imago; Solu Khumbu District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.727776&amp;materialsCitation.latitude=27.644722" title="Search Plazi for locations around (long 86.727776/lat 27.644722)">Paiya</a> [= Puiyan]; 27°38′41″ N, 86°43′40″ E (WGS 84); 8 Jun. 2003; Annemarie Ohler and Nicolas Pruvost leg.; MNHN 2003.3038 to MNHN 2003.3040  .</p><p>Other material examined</p><p>NEPAL • 4 tadpoles; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.333336&amp;materialsCitation.latitude=27.583334" title="Search Plazi for locations around (long 86.333336/lat 27.583334)">Ramechhap District</a>; 27°35′ N, 86°20′ E (WGS 84); 3084 m a.s.l.; Jun. 1997; local resident leg.; NHME (uncatalogued, field numbers 9705.10, 9705.15, 9705.21, 9705.22)  •  4 tadpoles; same locality as for holotype; Jun. 1997; local resident leg.; NHME (uncatalogued, field numbers 9705.73–76) •   3 tadpoles; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.23333&amp;materialsCitation.latitude=27.666666" title="Search Plazi for locations around (long 86.23333/lat 27.666666)">Dolakha District</a>; 27°40′ N, 86°14′ E (WGS 84); 2790 m a.s.l.; Jun. 1997; local resident leg.; NHME (uncatalogued, field numbers 9705.99, 9705.100, 9705.102)  .</p><p>Description (holotype)</p><p>Adult male, well preserved (Fig. 7). Measurements are provided in Table 3. SVL 41.6 mm.</p><p>HEAD. Large and flat, wider than long (HW /HL = 1.27); snout short and rounded; canthus rostralis distinct; nostril dorsolateral, just below canthal, midway between tip of snout and eye (NSD / END = 1.10); loreal region slightly concave; eye large (ED /HL = 0.43); internarial surface flat (IND = 4.20); pupil vertical; interorbital space flat; tympanum and tympanic ring absent; tongue oval; choanae small, visible when viewed from below; vomerine and maxillary teeth absent; vocal sac absent; supratympanic fold distinct, extending from posterior corner of eye to supra-axillary region, associated with parotoid glands.</p><p>FORELIMBS. Robust; forearm of median length (FAL / SVL = 0.29) and longer than hand (HAL / SVL = 0.25), without spines; fingers slender, free of dermal fringes or web; all fingertips rounded, not dilated; relative finger lengths: 3&gt; 4&gt; 2&gt; 1; subarticular tubercles absent; inner metacarpal tubercle flat and large; outer metacarpal tubercle indistinct; small black nuptial spines on dorsal and lateral surface of first and second fingers, and on inner side of third finger.</p><p>HINDLIMBS. Robust, moderately long (TIBL /SVL = 0.43; FEL / TIBL = 1.00), heels are not in contact when folded at right angles to the body (see Fig. 8 and Supp. file 1L); foot same length as shank; tips of toes round; toes not webbed, relative lengths 4&gt; 3&gt; 5&gt; 2&gt; 1; subarticular tubercles absent; moderately large inner metatarsal tubercle, outer metatarsal tubercle absent.</p><p>SKIN. Body dorsally and laterally with distinct tubercles in life and in preservative, each tubercle with one or two keratinized tips, only a few tubercles with spines; few scattered tubercles on upper and lower mandibles; forehead and surfaces of lower arm and tarsus relatively smooth; tubercles present below and on supratympanic fold; upper arms and legs with spineless tubercles; throat and belly surface smooth; a pair of pectoral glands and a pair of axillary glands present on chest, pectoral glands slightly larger than axillary, pectoral glands covered by small black spines.</p><p>Coloration</p><p>In life, dorsal surface of head, body, and extremities brownish; a small dark brown band extending from anterior edge of eye to nostril and further to tip of snout, and from posterior edge of eye along supratympanic fold; irregular dark brown spots or transverse stripes on all limbs including fingers and toes; ventral and dorsal surface of lower forelimbs and dorsal surface of hands with small irregular creamy white warts; flanks light brown; throat, belly, and partly ventral surfaces of extremities light melon-yellow; belly covered by irregular gray-brown network.</p><p>In preservative, color of dorsal surfaces has changed from brown to gray and that of ventral surfaces from yellowish to grayish white.</p><p>Variation</p><p>Measurements of the type series are provided in Table 3. Color can vary substantially from light to dark brown and even to olive (Supp. file 1L–M).</p><p>Distribution</p><p>Occurs in mountain forests in high-montane areas of the drainage area of the Koshi basin, Northeastern Nepal, more specifically, along the Khumbu Himal and adjacent Makalu region between ca 2800 and 3900 m a.s.l. (Fig. 1; Supp. file 1P). This species is known from the Sagarmatha and Koshi Zone, specifically the Dolakha District (Jiri), Ramechhap District, Solu Khumbu District (Lamjura La, Paiya, Surkie La, Taksindu), Bhojpur District (Salpa Pokhari), and in the Sankhuwasabha District. Eastern populations of this species that occur in the Bhojpur District (Salpa Pokhari), and in the Sankhuwasabha District are genetically distinct. This genetic differentiation supports the recognition of the eastern populations as a separate subspecies (see below).</p><p>Taxonomic remarks</p><p>In phylogenetic analyses, the new species belongs to a clade that branches with the clades previously identified in the genus  Scutiger . The species distinction is supported by substantial molecular divergence from other  Scutiger species, namely ≥ 11.1% uncorrected distance in the COI, ≥ 9.8% in the cytb, and ≥ 2.5% in the 16S gene (Supp. file 1D–F), by the resulting phylogenetic divergence (Fig. 2), as well as by morphological differences (Figs 4–5).  Scutiger khumbu sp. nov. features two clearly divergent, fully allopatric mitochondrial lineages: one from the Khumbu Himal and the other from the Makalu region. These lineages show high mitochondrial sequence divergence, with uncorrected distance in COI, cytb, and 16S ranging 8.5–9.3%, 5.0–7.4%, and 2.4–2.6% (Supp. file 1G–I), respectively, private alleles at the nuclear genes bfib7 and ccnb2, allele sharing at rag1, and lack notable morphological differences. Bioacoustic variation in the new species remains to be investigated. Given their deep phylogeographic structure, but younger divergence (including some nuclear allele sharing) and the absence of clear phenotypic differences, we consider the lineage from the Makalu region as distinct subspecies of  Scutiger khumbu sp. nov., namely  Scutiger khumbu makalu subsp. nov.</p><p>Remarks on larvae</p><p>Eleven tadpoles of western populations of  Scutiger khumbu sp. nov. at different stages ranging from 26 to 39 (Gosner 1960) (Fig. 9); LTRF: 3(2–3)/3(1–2), 3(2–3)/3(1–3), 3(2–3)/4(1–3), or 4(2–4)/4(1–3).</p></div>	https://treatment.plazi.org/id/05278791FFC6FFACEEDB04982D56FDCD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hofmann, Sylvia;Ohler, Annemarie;Baniya, Chitra B.;Dubois, Alain;Flecks, Morris;Jablonski, Daniel;Schmidt, Joachim;Dufresnes, Christophe	Hofmann, Sylvia, Ohler, Annemarie, Baniya, Chitra B., Dubois, Alain, Flecks, Morris, Jablonski, Daniel, Schmidt, Joachim, Dufresnes, Christophe (2025): Resolving taxonomic ambiguities in extant Himalayan lazy toads: A new species of the genus Scutiger (Anura, Megophryidae) from eastern Nepal. European Journal of Taxonomy 989 (1): 189-214, DOI: 10.5852/ejt.2025.989.2893, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2893/13117
05278791FFDCFFADEE6007ED2B76F902.text	05278791FFDCFFADEE6007ED2B76F902.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scutiger khumbu subsp. makalu Hofmann & Ohler & Baniya & Dubois & Flecks & Jablonski & Schmidt & Dufresnes 2025	<div><p>Scutiger khumbu makalu subsp. nov.</p><p>urn:lsid:zoobank.org:act: A5A7AE1C-43D3-4489-ACDA-AE0DEB942DB3</p><p>Figs 6, 9–10; Tables 1, 3; Supp. file 1J</p><p>Diagnosis</p><p>Scutiger khumbu makalu subsp. nov. can be distinguished from the nominotypical subspecies and other  Scutiger taxa based on multiple nucleotide substitutions in mitochondrial COI, cytb, and 16S sequences, as well as private alleles at the nuclear genes bfib7 and ccnb2 (Supp. file 1D–I), but not at the nuclear gene rag1. Specifically in the barcoding gene COI,  Scutiger khumbu makalu subsp. nov. is separated from the nominotypical subspecies and other currently recognized  Scutiger species by two binary and four asymmetric molecular signature characters (Table 1). Morphologically, this taxon closely resembles the nominotypical subspecies of  Scutiger khumbu sp. nov. but it exhibits a relatively larger foot length and tubercles that are less densely covered with small, keratinized tips, which also do not appear as dark as in the nominotypical subspecies (Fig. 6).</p><p>Etymology</p><p>The subspecific epithet ʻ makalu ʼ refers to the Makalu region in Nepal.</p><p>Type material</p><p>Holotype</p><p>NEPAL • ♂; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=86.93361&amp;materialsCitation.latitude=27.446388" title="Search Plazi for locations around (long 86.93361/lat 27.446388)">Bhojpur District</a>, Salpa Pokhari; 27°26′47″ N, 86°56′01″ E (WGS 84); 22 Jun. 2003; Thierry Deuve leg.; donated by Thierry Deuve; MNHN 2003.3041.</p><p>Paratype</p><p>NEPAL • 1 ♂; same locality as for holotype; 22 Jun. 2003; Thierry Deuve leg.; donated by Thierry Deuve; MNHN 2003.3042 .</p><p>Other specimens examined</p><p>NEPAL • 5 tadpoles; <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=87.21667&amp;materialsCitation.latitude=27.633333" title="Search Plazi for locations around (long 87.21667/lat 27.633333)">Sankhuwasabha District</a>; 27°38′ N, 87°13′ E (WGS 84); 3600 m a.s.l.; Jun. 1997; local residents leg.; NHME (uncatalogued, field numbers JS1405.23–27)  .</p><p>Description (holotype)</p><p>Adult male, well preserved (Fig. 10). Measurements are provided in Table 3. SVL = 47.9 mm.</p><p>HEAD. Large and flat, wider than long (HW/HL = 1.22); snout short and rounded; canthus rostralis distinct; nostril dorsolateral, just below canthal, midway between the tip of snout and eye (NSD/END = 1.00); loreal region slightly concave; eye relatively large (ED/HL = 0.37); internarial surface flat (IND = 5.10); pupil vertical; interorbital space flat; tympanum and tympanic ring absent; tongue oval; choanae small, visible when viewed from below; vomerine and maxillary teeth absent; vocal sac absent; supratympanic fold distinct, extending from posterior corner of eye to supra-axillary region, associated with parotoid glands.</p><p>FORELIMBS. Robust; forearm of median length (FAL/SVL = 0.28) and longer than hand (HAL/SVL = 0.26), without spines; fingers slender, free of dermal fringes or web; all fingertips rounded, not dilated; relative finger lengths: 3&gt; 4&gt; 2&gt; 1; subarticular tubercles absent; inner metacarpal tubercle flat and large; outer metacarpal tubercle large, nearly round, and flat; black nuptial spines on dorsal and lateral surface of first and second fingers, and on inner side of third finger.</p><p>HINDLIMBS. Robust, moderately long (TIBL/SVL = 0.41; FEL/TIBL = 1.10), heels are not in contact when folded at right angles to the body (see also Fig. 10); foot longer than shank; tips of toes round; toes not webbed, relative lengths 4&gt; 3&gt; 5&gt; 2&gt; 1; subarticular tubercles absent; moderately large inner metatarsal tubercle, outer metatarsal tubercle absent.</p><p>SKIN. Body dorsally and laterally with distinct tubercles in preservative, each tubercle with one large keratinized spine in the center (Fig. 6B), surrounded by small, keratinized dark brown or blackish tips; small scattered spines on upper and lower mandibles; dorsal surface of forehead and surfaces of lower arm relatively smooth; tubercles with spines present below and on supratympanic fold; upper arms and legs with tubercles, most of them with spines; throat and belly surface smooth; a pair of pectoral glands and a pair of axillary glands present on chest, connected, pectoral glands larger than axillary, all covered by small black spines.</p><p>Coloration</p><p>In preservative, dorsal surfaces gray, ventral surfaces grayish white.</p><p>Distribution</p><p>This subspecies is known from cloud forests in high-montane areas in the Makalu region in Nepal at altitudes between ca 3000 and 3600 m a.s.l., specifically in the Bhojpur District (Salpa Pokhari), and in the Sankhuwasabha district.</p><p>Taxonomic remarks</p><p>Scutiger khumbu makalu subsp. nov. was previously identified as a distinct phylogeographic lineage by Hofmann et al. (2017) based on molecular data of a few samples. It is characterized by high genetic divergence in the COI (8.8%), cytb (6.3%), and 16S (2.5%) genes, distinct alleles at the nuclear bfib7 and ccnb2 genes, and shared alleles at the rag-1 gene, with the nominotypical subspecies of  Scutiger khumbu sp. nov., to which it is allopatric and lacks a clear morphological differentiation. The divergence being lower than in most  Scutiger species, it is therefore described as a new subspecies of  Scutiger khumbu sp. nov.</p><p>Remarks on larvae</p><p>Five tadpoles of  Scutiger khumbu makalu subsp. nov. at different stages ranging from 26 to 29 (Gosner 1960) (Fig. 9); LTRF: 3(2–3)/3(1–2) or 3(2–3)/4(1–3).</p></div>	https://treatment.plazi.org/id/05278791FFDCFFADEE6007ED2B76F902	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Hofmann, Sylvia;Ohler, Annemarie;Baniya, Chitra B.;Dubois, Alain;Flecks, Morris;Jablonski, Daniel;Schmidt, Joachim;Dufresnes, Christophe	Hofmann, Sylvia, Ohler, Annemarie, Baniya, Chitra B., Dubois, Alain, Flecks, Morris, Jablonski, Daniel, Schmidt, Joachim, Dufresnes, Christophe (2025): Resolving taxonomic ambiguities in extant Himalayan lazy toads: A new species of the genus Scutiger (Anura, Megophryidae) from eastern Nepal. European Journal of Taxonomy 989 (1): 189-214, DOI: 10.5852/ejt.2025.989.2893, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2893/13117
