identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
62199F50FFECFFC3FF34FCC3FCFFF8BC.text	62199F50FFECFFC3FF34FCC3FCFFF8BC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes palavanensis (Boulenger 1894)	<div><p>Limnonectes palavanensis (Boulenger, 1894)</p><p>Rana palavanensis — Boulenger 1894: 85. Type locality: “ Palawan ”; Boulenger 1897: 230 (partim); Boulenger 1920: 59 (partim); van Kampen 1923: 182 (partim).</p><p>Rana microdisca palavanensis — Inger 1954: 299 (partim); Inger 1966: 222 (partim).</p><p>Limnonectes palavanensis — Setiadi et al. 2011: 224; Inger et al. 2017: 87 (partim).</p><p>Lectotype (by present designation). BMNH 1947.2.1.96, adult female, from Palawan, collected by A. Everett (Fig. 3A).</p><p>Paralectotypes (by present designation). BMNH 1947.2.1.94, 1947.2.1.95, two adult females, from Palawan, collected by A. Everett.</p><p>Referred specimens. KU 309130, adult male, from Mt. Mantalingahan (8.7637167°, 117.6867°; 650 m), Barangay Samarinana, Municipality of Brooke’s Point, Palawan Province, Palawan Island, Philippines . KU 309129, from Irawan Watershed (9.8368°, 118.64125°; 180 m), Barangay Irawan, Municipality of Puerto Princesa, Palawan Province, Philippines .</p><p>Diagnosis. Limnonectes palavanensis belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing moderately extensive; supratympanic fold curved; pineal spot weakly discernible, at level of anterior upper-eyelid edge; vomer ridges long, narrowly separated from each other; postpalpebral ridges and lumbar ridges low, indistinct and interrupted; median ridge absent; size small with SVL of adult females 31.6–32.3 mm. The species differs from all other species of the group by at least 6.2% in the sequence of the 16S barcoding gene.</p><p>Description of lectotype. Adult female; SVL 31.6 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3A); head large (HL/SVL 0.36, HW/SVL 0.35), about as wide as long (HW/HL 0.97); snout moderately long (SL/HL 0.43), subacuminate in dorsal view, truncate and slightly protruding in lateral profile, wider than long (SL/EE 0.85); canthus rostralis moderately distinct between eye and nostril, slightly concave from eye to nostril in dorsal view; loreal region oblique; nostrils rounded, directed dorsolaterally, situated about halfway between tip of snout and eye (EN/NS 0.97), separated from each other by distance much larger than distance between eye and nostril (NN/EN 1.56); eye directed anterolaterally, moderately protruding, large (ED/HL 0.33), its diameter shorter than snout (ED/SL 0.76); interorbital distance greater than upper eyelid width (IO/EW 1.22) and smaller than internarial distance (IO/NN 0.74); pineal spot weakly discernible, at level of anterior upper-eyelid edge; tympanum and its annulus distinctly and fully visible, not covered by supratympanic fold; tympanum separated from eye by little less than one-third its diameter (ET/TD 0.31); tympanum diameter smaller than eye diameter (TD/ED 0.57); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, lateral fifth covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, long, separated from choana by about two-thirds length of individual process and from each other by half length of individual process (Fig. 4A); tongue moderately long and narrow, bilobed for about oneeighth its length, free distally for about two-fifths its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs and lateral surfaces of trunk finely shagreened; small tubercles sparsely scattered on upper eyelid and on dorsum; dorsolateral dermal ridge distinct on both sides from posterior end of upper eyelid to groin; postpalpebral ridge formed by small tubercles, low, interrupted and indistinct; horseshoe-shaped dermal ridge low, in middle of back just behind level of arms; lumbar ridge indistinct, low and interrupted, from near posterior end of horseshoe-shaped ridge to supraanal region; median ridge absent; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, curved (Fig. 5A); ventral side of head smooth; transverse chest fold between arm insertions low; ventral side of trunk weakly areolate; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: I = II = IV &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, prominent, about one-third length of metacarpal of Finger I; inner and outer palmar tubercles fused proximally, forming large, roughly U-shaped, prominent tubercle on proximal half of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.86); heel reaching half SL beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.63), longer than thigh (TFL/THL 1.17); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot shorter than tibiofibula (FOT/TFL 0.82); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1+/2 II 1+/2.25 III 1.5/3- IV 3-/1.5 V (Fig. 6); narrow dermal ridge on preaxial side of Toe I and on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. (from Boulenger 1894:) Dorsum brown; sides of snout below canthus blackish; temporal region light; dark cross bar between eyes; horseshoe-shaped tubercle blackish; dorsolateral folds edged with blackish laterally; limbs with regular dark cross bands; ventral side whitish. See Fig. 7A for a photograph of a specimen in life.</p><p>In preserved state (Fig. 3A), dorsum of lectotype including flaks and dorsal parts of limbs light brown; lower arm, fingers, legs, and feet with mid-brown crossbars; upper eyelids with bluish tinge; interorbital bar mid-brown; supratympanic fold and dorsolateral fold as well as area between these folds mid-brown; tympanum cream-coloured with light brown rim; sides of head mid-brown; two mid-brown blotches on dorsal side of snout; horseshoe-shaped tubercle dark brown; ventral side of head, chest, arms and legs yellowish cream-white; abdomen whitish cream-coloured; infralabial region dark brown with small white spots; throat without pattern.</p><p>Variation. The female paralectotypes match the holotype in general appearance.The state of BMNH 1947.2.1.94 does not allow it to be measured. SVL of BMNH 1947.2.1.95 is 32.3 mm. It matches the lectotype in proportions and measurements (Table 2). Pedal webbing variation is I 1+/2 II 1+/2.25 [50], 2.5 [50] III 1.5/3- IV 3-/1.5 V (Fig. 6). Boulenger (1894, 1897) stated an SVL of 43 mm for the species, later corrected to 31–33 mm for the three type specimens (Boulenger 1920).</p><p>Bioacoustics. The advertisement call and other vocalisations are unknown.</p><p>Distribution. The species is restricted to Palawan, Philippines where it occurs at elevations between 180 m and 650 m.</p><p>Ecology. Limnonectes palavanensis is found in swamps, seepages, and wide, slow-moving streams (Setiadi et al. 2011). Alcala &amp; Brown (1998) treat the species as Rana microdisca in an account together with Limnonectes parvus from Mindanao, Philippines, and the ecological data provided in that account most likely refer to the latter.</p><p>Etymology. Named after the type locality, Palawan Island in the Philippines.</p><p>Suggested English name. Palawan Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFECFFC3FF34FCC3FCFFF8BC	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFF5FFDEFF34FF50FCE1F8FC.text	62199F50FFF5FFDEFF34FF50FCE1F8FC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes gunungensis Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes gunungensis sp. nov.</p><p>Holotype. NMBE 1061576, adult female, from Pa’ Lungan (3.808622°, 115.524576°; 1,100 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia, collected on 5 March 2012 by S. T. Hertwig, A. Jankowski and Y. M. Pui (Fig. 3B).</p><p>Paratypes. NMBE 1061593, juvenile, from the type locality, collected on 4 March 2012 by S. T. Hertwig, A. Jankowski and Y. M. Pui; NMBE 1065994, adult female, from Payeh Maga (4.4°, 115.5°), Sarawak, collected on 2 March 2013 by S. T. Hertwig, Y. M. Pui et al .; NMBE 1073882, adult female, from “Old camp”, Payeh Maga, collected on 28 February 2016; NMBE 1073867, adult male, NMBE 1073886–88, 1073891, four adult females, from a stream behind Camp 1 (4.438°, 115.538°; 960 m), Payeh Maga, Sarawak, Malaysia, collected on 29 February 2016 ; all collected by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen . NMBE 1101226, collected on 25 February 2019; NMBE 1101230, collected on 26 February 2019; NMBE 1101231, collected on 27 February 2019, three adult males, from Kerangas Forest, Camp Balah (3.127907°, 114.278603°; 1,074 m), Gunung Dulit, Sarawak, Malaysia, collected by L. Etter, S . T. Hertwig and F. Heussler . NMBE 1101220–1101221, 1101232, three adult males, from Camp Koyan (3.342366°, 114.177573°; 822 m), High Plateau, Gunung Dulit, Sarawak, Malaysia, collected on 5 March 2019 by L. Etter, S. T. Hertwig and F. Heussler .</p><p>Referred specimens. BMNH 1978.213, adult female, from Camp 3, Gunung Mulu, Gunung Mulu National Park, Sarawak, Malaysia, collected on 7–8 September 1977 by J. Dring ; NMBE 1101227–1101229, three tadpoles from Kerangas Forest, Camp Balah (3.127907°, 114.278603°; 1,074 m), Gunung Dulit, Sarawak, Malaysia, collected with an adult male paratype (NMBE 1101226) .</p><p>Diagnosis. Limnonectes gunungensis sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing extensive; supratympanic fold subangular; pineal spot well discernible, at level of anterior upper-eyelid edge; vomer ridges long, narrowly separated from each other; postpalpebral ridge moderately distinct, along anterior part of dorsolateral ridge to just behind level of arms; lumbar and median ridges absent; size relatively large with SVL of adult males 25.6–31.5 mm, of adult females 30.4–39.9 mm. The species differs from all other species of the group by at least 6.7% in the sequence of the 16S barcoding gene. The species occurs at mid-level elevations (1,000–1,600 m) in the transition between submontane rainforest and lower montane rainforest .</p><p>Description of holotype. Adult female; SVL 38.6 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3B); head large (HL/SVL 0.37, HW/SVL 0.38), about as wide as long (HW/HL 1.01); snout moderately long (SL/HL 0.43), subacuminate in dorsal view, rounded and slightly protruding in lateral profile, as long as wide (SL/EE 0.98); canthus rostralis weakly expressed between eye and nostril, slightly concave from eye to nostril in dorsal view; loreal region oblique; nostrils rounded, directed laterally, situated about halfway between tip of snout and eye (EN/NS 1.08), separated from each other by distance much larger than distance between eye and nostril (NN/EN 1.34); eye directed anterolaterally, moderately protruding, large (ED/HL 0.32), its diameter shorter than snout (ED/SL 0.74); interorbital distance greater than upper eyelid width (IO/EW 1.23) and smaller than internarial distance (IO/NN 0.83); pineal spot well discernible, at level of anterior upper-eyelid edge; tympanum and its annulus distinctly and fully visible, its posterior fifth obscured by thickened skin, its dorsal edge in contact with supratympanic fold; tympanum separated from eye by about half its diameter (ET/TD 0.53); tympanum diameter slightly more than half eye diameter (TD/ED 0.55); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, lateral fifth covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, long, separated from choana by distance about length of individual process and from each other by half length of individual process (Fig. 4B); tongue moderately long and broad, bilobed for about one-sixth its length, free distally for about three-fourth its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs and lateral surfaces of trunk finely shagreened; dorsolateral ridge prominent, from shortly behind posterior end of upper eyelid to groin; postpalpebral ridge moderately distinct, only in anterior part of dorsum, from behind eyes to just behind level of arms; lumbar and median ridges absent; horseshoe-shaped dermal ridge low, indistinct, just behind level of arms; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, subangular, with anterior and posterior part about equal in length (Fig. 5B); ventral side of head smooth; weak transverse chest fold between arm insertions; ventral side of trunk weakly areolate; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.26); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: II &lt;I = IV &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, prominent, about one-third length of metacarpal of Finger I; inner palmar tubercle subcircular, about as long as narrow outer palmar tubercle; palmar tubercles fused proximally on proximal half of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.85); heel reaching slightly beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.59), longer than thigh (TFL/THL 1.12); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.94); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1/2 II 1+/2+ III 1.25/2.5 IV 2.5/1+ V (Fig. 6); narrow dermal ridge on preaxial side of Toe I and on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life, dorsum dark brown with narrow, whitish median band; limbs including fingers and toes somewhat lighter brown with regular medium brown crossbands; interorbital bar dark brown, interrupted by light median band; median band edged with dark brown on dorsal part of snout; dorsolateral fold blackish; horseshoe-shaped tubercle black laterally; fine line of black dots and speckling from posterior ends of horseshoe-shaped tubercle along median band to vent, much wider and conspicuous in posterior part; ventral side of head and chest cream-coloured with an orange tinge; limbs reddish orange; abdomen light yellow. See Fig. 7B for a photograph of a male paratype in life.</p><p>In preserved state, dorsum dark brown with narrow, whitish median band (Fig. 3B); crossbands on limbs, fingers and toes dark brown; interorbital bar black; median band edged with black on dorsal part of snout; dorsolateral fold blackish; horseshoe-shaped tubercle black laterally; fine line of black dots and speckling from posterior ends of horseshoe-shaped tubercle along median band to vent, much wider and conspicuous in posterior part; ventral side of head and chest cream-white; limbs yellowish, abdomen hardly pigmented, blueish white.</p><p>Variation. The paratypes match the holotype in general appearance. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally,situated at corner of mouth.SVLof adult males is 25.6–31.5 mm,of adult females 30.4–39.9mm (Table 2). Pedal webbing variation is I 1 [40],1+ [60] /2 [90],2+ [10] II 1 +/2+ [20],2.25 [60],2.5 [20] III 1 + [10],1.25 [70],1.5 [20] /2.5 [30],2.75 [30],3- [40] IV 2.5 [10],2.75 [30],3- [20], 3 [40] /1+ [50],1.25 [50] V (Fig. 6). The median dorsal band is absent in most specimens .</p><p>Bioacoustics. The advertisement call and other vocalisations are unknown, but see account on bioacoustics of L. nephophilus sp. nov. below.</p><p>Distribution. The species is so far known from only a few places in eastern Sarawak between 1,000 and 1,600 m (Figs. 8, 9). Apart from the type locality, Pa’ Lungan, it has been recorded from other locations around Gunung Murud and from Payeh Maga, Gunung Dulit, and Gunung Mulu (Fig. 8). The species co-occurs sympatrically with L. sarawakensis sp. nov. on the High Plateau on Gunung Dulit, Sarawak, and with L. nephophilus sp. nov. around Camp 3 on Gunung Mulu, Sarawak (Fig. 8).</p><p>Ecology. Adult males and gravid females were collected in moist leaf litter close to small streams. One of the male paratypes (NMBE 1101226) was carrying tadpoles (NMBE 1101227–1101229) on its back (Fig. 10E). The number and size of the ripe ova of the females indicate small clutch sizes .</p><p>Etymology. The species epithet refers to the distribution of the species on mountains in Sarawak (including Gunung Murud, Gunung Dulit, and Gunung Mulu) and is composed of the Malay word “gunung” (meaning “mountain”) as an invariable noun and the Latin suffix -ensis (“from”); meaning “inhabitant of the mountain”.</p><p>Suggested English name. Mountain Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFF5FFDEFF34FF50FCE1F8FC	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFF3FFDBFF34FA9FFCE7F818.text	62199F50FFF3FFDBFF34FA9FFCE7F818.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes kinabaluensis Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes kinabaluensis sp. nov.</p><p>Rana palavanensis (nec Rana palavanensis Boulenger, 1894): Boulenger 1897: 230 (partim); Boulenger 1920: 59 (partim); van</p><p>Kampen 1923: 182 (partim); Smith 1931: 30; Inger &amp; Stuebing 1992: 46 (partim); Hoffman 1994: 223, 1995: 11; Inger et al. 1996: 363; Lakim et al. 2000: 32. Rana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1966: 222 (partim); Matsui 1979: 333. Hylarana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894): Manthey 1983: 22. Limnonectes palavanensis (nec Rana palavanensis Boulenger, 1894): Malkmus 1994: 244; Hoffmann 2002: 31; Malkmus et al.</p><p>2002: 145 (partim).</p><p>Holotype. NMBE 1075310, adult female, from Silau-Silau Trail (6.012741°, 116.539129°, 1,604 m), Gunung Kinabalu, Kinabalu National Park, Sabah, Malaysia, collected on 5 March 2017 by A. Haas, R. Hagmann, S. T. Hertwig and P. Yambun (Fig. 3C).</p><p>Paratypes. SP 2076, adult male, from Silau-Silau Trail, Headquarters, Kinabalu National Park, Sabah, Malaysia, collected on 9April 1997 by “Paul, RFI” [= Paul Yambun and Robert F. Inger]. SP 2579, 2582–2584, one female, two males, one subadult, from Bundu Tuhan Trail, Headquarters, Kinabalu National Park, Sabah, Malaysia, collected on 23 December 1999 by “Maklarin” [bin Lakim], “Johny”, “Rashid”, David” [Sumpongul] and “Ben”[edict Butit] .</p><p>Referred material. ZFMK 79834, adult male, from Sungai Wariu, Sayap, Sabah, collected on 16 June 1993 by P. Hoffmann. SP 1296, subadult, collected on 14 June 1995 by “Maklarin [bin Lakim], Alim et al.”; SP 1355, adult female, collected on 15 June by the Zoologi Unit of Sabah Parks; both from Kota Belud, Sayap Park Substation, Kinabalu National Park, Sabah, Malaysia. SP 1395, adult female, collected in July 1995 by the Zoologi Unit; SP 1545, juvenile, both from Sayap, Kinabalu National Park, Sabah, Malaysia. SP 1769, subadult female, collected on 23 March by the Zoologi Unit; SP 27184, juvenile, collected on 10 October 2011 by “Paul” [Yambun], “Martinah” [Latim], “Dominik” [Tinggoh]; both from Lipasu Trail, Sayap Park Substation, Kinabalu National Park, Sabah, Malaysia. SP 558, adult of undetermined sex, from “Sungai (Bekalan Air)”, Poring, Kinabalu Park, Sabah, collected on 27 September 1980 by R. Stuebing. SP 2864, adult male, from Gunung Trusmadi, collected on 1 November 2001 by “Maklarim etc.” . UNIMAS 9061, adult female, from Lipasu Trail, Sayap, Sabah, Malaysia, collected on 23 August 2007 by I. Das, A. Haas, K. B. Hee and C. Garrett . NMBE 1072214, adult female, collected on 14 March 2015; NMBE 1072159, juvenile, collected on 17 March 2015; both collected by S . T. Hertwig et al. from Mountain Hostel (4.4611°, 117.9230°; 900 m), Tawau Hills National Park, Sabah, Malaysia .</p><p>Diagnosis. Limnonectes kinabaluensis sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing reduced; supratympanic fold subangular with its anterior part and posterior part about equal in length; pineal spot well discernible, at level of anterior upper-eyelid edge; vomer ridges short, very widely separated from each other; median ridge low, glandular, moderately distinct, only from horseshoe-shaped ridge to cloaca; postpalpebral and lumbar ridges absent; size relatively large with SVL of adult males 29.3–31.8 mm, of adult females 32.4–38.0 mm. The species differs from all other species of the group by at least 5.7% in the sequence of the 16S barcoding gene. It occurs in submontane and montane rainforest at elevations between 850 and 1,604 m.</p><p>Description of holotype. Adult female; SVL 35.5 mm; body sturdy, widest at temporal region, slightly tapering to groin (Fig. 3C); head large (HL/SVL 0.38, HW/SVL 0.40), wider as long (HW/HL 1.06); snout moderately long (SL/HL 0.43), rounded in both dorsal view and lateral profile, slightly wider than long (SL/EE 0.95); canthus rostralis distinct between eye and nostril, weakly concave in dorsal view; loreal region oblique; nostrils rounded, directed laterally; situated slightly closer to tip of snout than to eye (EN/NS 1.06), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.44); eye directed anterolaterally, moderately protruding, moderately large (ED/HL 0.33), its diameter much shorter than snout (ED/SL 0.76); interorbital distance greater than upper eyelid width (IO/EW 1.48) and smaller than internarial distance (IO/NN 0.83); pineal spot well discernible, at level of anterior upper-eyelid edge; tympanum and its annulus distinctly visible, its posterior fifth covered by thick skin; tympanum separated from eye by little less than half its diameter (ET/TD 0.44); tympanum diameter smaller than eye diameter (TD/ED 0.63); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, subtriangular, located far anterolaterally at margins of roof of mouth, fully visible in ventral view; vomer processes bearing teeth, short, separated from choana by distance about equal to length of individual process, from each other by 1.5 times length of individual process (Fig. 4C); tongue moderately long and narrow, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs finely shagreened; lateral surfaces of trunk finely areolate with sparsely scattered small tubercles; dorsolateral ridge prominent, from posterior end of upper eyelid to groin; low, horseshoe-shaped dermal ridge distinct, in middle of back just behind level of arms; median ridge low, glandular, moderately distinct, only on posterior part of dorsum, from horseshoe-shaped ridge to cloaca; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, subangular with its anterior part and posterior part about equal in length (Fig. 5C); enlarged tubercles on lateral edge of upper eyelid; ventral side of head smooth with several weak wrinkles at lateral edge near mandible; transverse chest fold between arm insertions weakly distinct; ventral side of trunk weakly areolate, almost smooth; ventral side of limbs smooth; cloaca without dermal flap but with densely scattered small tubercles.</p><p>Forelimbs sturdy; hand relatively small (HND/SVL 0.26); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: IV &lt;II &lt;I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, flat, about two-fifths length of metacarpal of Finger I; inner palmar tubercle rounded, weakly distinct; outer palmar tubercle elongate, about half length of metacarpal, moderately distinct.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.84); heel reaching beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL0.60), longer than thigh (TFL/THL1.15); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.88); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1.5/2+ II 1.25/3- III 2/3.25 IV 3.25/2 V (Fig. 6); narrow dermal flap on preaxial side of Toe I between metatarsal tubercle and disk and on postaxial side of Toe V, widest from proximal end of metacarpus to proximal subarticular tubercle, continued to disk; inner metatarsal tubercle very prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life, (Fig. 7C), dorsum, limbs including fingers and toes light brown; upper arms without crossbands; lower arms, fingers, legs including feet with regular medium brown crossbands; interorbital bar dark brown; supratympanic fold and dorsolateral fold greyish dark brown; tympanum light brown; large light brown spots below eye on each side; horseshoe-shaped tubercle same colour as dorsum except dark brown posterior edge; four large dots arranged in rhombic pattern at level of pelvis, dark brown, each with large light grey tubercle in centre; area around cloaca black with whitish small tubercles; ventral side of head, chest, arms and legs orange; large dark brown spots in infralabial region; throat speckled with minute cream-coloured dots; abdomen yellow.</p><p>In preserved state (Fig. 3C), dorsum greyish brown; limbs, fingers and toes light brown; regular crossbands on lower arms, fingers, legs, feet dark brown; interorbital bar dark brown; supratympanic fold and dorsolateral fold blackish; tympanum dark brown; large spots below eye dark brown; black posterior edge of horseshoe-shaped tubercle black; four large dots arranged in rhombic pattern at level of pelvis dark brown, each with large grey tubercle in centre; area around cloaca black with whitish small tubercles; ventral side of head, chest, arms and legs yellowish cream-white; large spots in infralabial region dark brown; throat speckled with minute dots; abdomen hardly pigmented, blueish white.</p><p>Variation. The paratypes match the holotype in general appearance and proportions. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. SVL of paratypes is 30.2–31.8 mm in adult males and 36.8 mm in the adult female (Table 2). Pedal webbing variation is I 1.5 [100] /2- [33],2 [33],2+ [33] II 1+ [66],1.25 [33] /2.5 [33],2.75 [33],3- [33] III 1.5 [66],2 [33] /3 [66],3.25 [33] IV 3 [66],3.25 [33] /1.5 [33],2- [33], 2 [33] V (Fig. 6). SVL of referred specimens from other localities is 29.3 mm in adult males and 32.4–38.0 mm in adult females.</p><p>Bioacoustics. The advertisement call of the species is unknown. The call description, including an audiospectrogram and a waveform, in Malkmus et al. (2002: 147, Fig. LXIV) is not based on a call of a specimen from Gunung Kinabalu but on the call recorded by J. Dring from Camp 3 in Gunung Mulu National Park that we refer to L. nephophilus sp. nov. (see below).</p><p>Distribution. The species is distributed around the headquarters of Kinabalu National Park Headquarters (Silau-Silau Trail, Bundu Tuhan View Trail) at about 1,600 m. Referred specimens have been collected at Sayap (850–900 m) on the western slope of Gunung Kinabalu; and we tentatively assign a badly preserved specimen (SP 558) from Poring on the eastern slope of Gunung Kinabalu and a specimen from Gunung Trusmadi (SP 2864) to the species. The species has a disjunct distribution and has also been collected at higher elevations of about 900 m in Tawau Hills National Park, Sabah, Malaysia, where it occurs parapatrically with Limnonectes phylax sp. nov. that is distributed at lower elevations in Tawau Hills National Park (Figs. 8, 9).</p><p>Ecology. The types were collected in leaf litter on the forest floor. The specimens from Sayap were collected under a log, another one on a rock, others on the ground and under leaves in leaf litter; both adults and juveniles were found on the ground in moist leaf litter in some distance from the nearest stream (Hoffmann 1994). According to Malkmus et al. (2002), adults of the species feed on ants, termites, and other small terrestrial invertebrates. The number and size of the ripe ova of the females indicate small clutch sizes. A male was observed releasing tadpoles from its back into a shallow puddle on Silau-Silau Trail in the Headquarters of Kinabalu National Park on 7 July 2023 (P. Janzen, in litt., Fig. 10F).</p><p>Etymology: The species epithet is composed of the name of the type locality, Gunung Kinabalu, as an invariable noun and the Latin suffix -ensis (“from”); meaning “an inhabitant of Kinabalu”.</p><p>Suggested English name. Kinabalu Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFF3FFDBFF34FA9FFCE7F818	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFF0FFD4FF34FC09FCCAF87A.text	62199F50FFF0FFD4FF34FC09FCCAF87A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes nephophilus Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes nephophilus sp. nov.</p><p>Rana palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1985: 52 (partim).</p><p>Holotype. NMBE 1056394, adult male, from Camp 3 (4.038079°, 114.888078°; 1,350 m), Gunung Mulu National Park, Sarawak, Malaysia, collected in March 2009 by J. M. Dehling, N. W. Tinggang, Y. M. Pui, S. T. Hertwig and A. Jankowski (Fig. 3D).</p><p>Paratypes. BMNH 1978.214 – 215, two adult males, collected on 7–8 September 1977 at the type locality by J. Dring; BMNH 1978.1666 – 1667, two adult females, BMNH 1978.1668 – 1669, two juveniles, all from the type locality, collected in 1978 by J. Dring . NMBE 1065176–1065177, two tadpoles, from Camp 4 (4.044178°, 114.913847°; 1,800 m), Gunung Mulu National Park, Sarawak, Malaysia , collected on 29 March 2007 by S. T. Hertwig and K. Lilje . NMBE 1061685, adult female, NMBE 1061686, adult male, collected on 24 February 2013 by S . T. Hertwig, A. Jankowski, Y. M. Pui; NMBE 1073768, adult male, NMBE 1073755, adult female, collected on 6 March 2016 by J. Flury, A. Haas, S . T. Hertwig, Y. M. Pui and N. Reichen; all from Lepo Bunga (3.941808°, 115.544667°; 1,760 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia . NMBE 1061641–1061642, two juveniles, collected on 29 February 2012 by S . T. Hertwig, A. Jankowski and Y. M. Pui; NMBE 1073581, adult male, NMBE 1073590, adult female, NMBE 1073706, 1073709, two juveniles, collected on 9–12 March 2016 by J. Flury, A. Haas, S . T. Hertwig, Y. M. Pui and N. Reichen; all from Church Camp (3.926317°, 115.513889°; 2,100 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia . NMBE 1073578, adult male, from Summit Trail (2,100 m), Church Camp, Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia , collected on 10 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen . NMBE 1073715, unsexed adult, from Lepo Bunga (3.941808°, 115.544667°; 1,760 m), Gunung Murud, Pulong Tau National Park, Sarawak, Malaysia , collected on 7 March 2016 by J. Flury, A. Haas, S. T. Hertwig, Y. M. Pui and N. Reichen .</p><p>Referred material. BMNH 1978.216, tadpole, from the type locality, collected on 8 September 1977 by Julian Dring .</p><p>Diagnosis. Limnonectes nephophilus sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing very reduced; supratympanic fold subangular with anterior part shorter than posterior part; pineal spot distinctly discernible, slightly anterior to level of anterior upper-eyelid edge; vomer ridges short, widely separated from each other; median ridge low, indistinct, running from level of nostrils to level of pelvis; postpalpebral and lumbar ridges absent; size relatively large with SVL of adult males 26.1–31.9 mm, of adult females 33.1–37.6 mm. The species differs from all other species of the group by at least 5.7% in the sequence of the 16S barcoding gene. The advertisement call is a sharp chirp, 75–90 ms in duration at a dominant frequency of 2.5–3.0 kHz with a marked frequency modulation, given in call bouts of up to 21 calls with an intercall interval of 135–383 ms. The species occurs in montane rainforest between 1,200 and 2,100 m.</p><p>Description of holotype. Adult male; SVL 28.4 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3D); head large (HL/SVL 0.37, HW/SVL 0.38), about as wide as long (HW/HL 1.03); snout moderately long (SL/HL 0.40), rounded in both dorsal view and profile, slightly projecting beyond lower jaw, wider than long (SL/EE 0.82); canthus rostralis distinct between eye and nostril, slightly concave; loreal region oblique, moderately concave; nostrils rounded, directed laterally, situated slightly closer to tip of snout than to eye (EN/NS 1.09), separated from each other by distance much larger than distance between eye and nostril (NN/EN 1.70); eye directed anterolaterally, moderately protruding, large (ED/HL 0.32), its diameter shorter than snout (ED/SL 0.80); interorbital distance greater than upper eyelid width (IO/EW 1.19) and smaller than internarial distance (IO/NN 0.69); pineal spot distinctly discernible, slightly anterior to level of anterior upper-eyelid edge; tympanum and its annulus distinctly visible, except posterior one-fourth which is covered by thick skin; tympanum separated from eye by little more than one-third of its diameter (ET/TD 0.37); tympanum diameter smaller than eye diameter (TD/ ED 0.61); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, short, separated from each other and from choana by length of individual process (Fig. 4D); tongue moderately long and narrow, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent; vocal sac apertures slit-like, directed posterolaterally, situated closer to corner of mouth than to base of tongue.</p><p>Dorsal surfaces of head, trunk and limbs finely shagreened; lateral surfaces of trunk finely areolate with scattered small tubercles; dorsolateral ridge prominent, from posterior end of upper eyelid to groin; median ridge low, indistinct, running from level of nostrils to level of pelvis; postpalpebral and lumbar ridges absent; horseshoe-shaped dermal ridge low, in middle of back just behind level of arms; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, subangular with anterior part shorter than posteroventrally directed posterior part (Fig. 5D); row of small tubercles on lateral edge of upper eyelid, postaxial side of forearm, and lateral part of head; ventral side of head smooth with weak dermal folds parallel to posterior part of mandible; weak transverse chest fold between arm insertions; ventral side of trunk weakly areolate with sparsely scattered small tubercles; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: II = IV &lt;I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, flat, little more than one-third length of metacarpal of Finger I; inner palmar tubercle indistinct, small, rounded, flat, on proximal end of metacarpus of Fingers II and III; outer palmar tubercle indistinct, flat, elongated, on proximal half of metacarpal of Finger IV; nuptial pads absent.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.84); heel reaching to tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.57), slightly longer than thigh (TFL/THL 1.11); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.93); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, slightly enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1.5/2.25 II 1.5/3- III 2.25/3.25 IV 3.25/2+ V (Fig. 6); dermal flap on postaxial side of Toe V, widest from proximal end of metacarpus to proximal subarticular tubercle, continued to disk; inner metatarsal tubercle very prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent; low ridges of callous tissue from proximal plantar part of metatarsus to proximal subarticular tubercle on Toes I, II, and V.</p><p>Colouration. In life (Fig. 7D), dorsum of holotype including lateral parts light brown with orange tinge; interorbital bar blackish brown; head anterior to interorbital bar as well as tympanic area bright orange; horseshoe-shaped tubercle slightly darker brown than dorsum and dark brown in the centre; two large black dots in longitudinal row at level of pelvis; area around cloaca dark brown; supratympanic fold dark grey; dorsolateral fold red in anterior part, lightening to orange posteriorly, bordered laterally by thin black line; tympanum light brown; side of snout light brown with larger dark brown spots and dark bar along canthus rostralis; arms orange, darkening to light brown towards hands; lower arm with dark brown crossbands; hands yellowish brown; legs reddish light brown with greyish light brown crossbands; tarsus and toes red; ventral side of head, chest, arms, lower legs and tarsus yellowish bright orange; large dark brown spots in infralabial region; throat speckled with minute brown dots; abdomen bright yellow, darkening laterally, with white speckling; thighs dark orange, darkening to reddish preaxially and postaxially; preaxial side of thighs with bluish white speckling; plantar side of feet blackish.</p><p>In preserved state (Fig. 3D), dorsum creamish light brown; arms, fingers and toes as well as dorsal part of snout yellowish light brown; upper eyelids light blue; legs reddish light brown; upper arms without crossbands; lower arms, fingers, legs including feet with regular dark brown crossbands; interorbital bar blackish brown; supratympanic fold greyish brown; dorsolateral fold bordered laterally by thin black line; tympanum light brown; three large dark brown spots below eye and nostril on each side; horseshoe-shaped tubercle slightly darker brown than dorsum and dark brown in the centre; two large black dots in longitudinal row at level of pelvis; area around cloaca dark brown; ventral side of head, chest, arms and lower legs yellowish cream-white; large dark brown spots in infralabial region; throat speckled with minute brown dots; abdomen hardly pigmented, blueish white; thighs reddish cream-coloured.</p><p>Variation. The paratypes match the holotype in general appearance. SVL of adult males is 26.1–31.9 mm, of adult females 33.1–37.6 mm (Table 2). Pedal webbing variation is I 1.5 [30],1.75 [40],2- [30] /2+ [20],2.25 [70],2.5 [10] II 1.5 /2.75 [10],3- [80],3 [10] III 2 [10],2+ [50],2.25 [40] /3.25 [80],3.5 [20] IV 3.25 [80],3.5 [20] /2 [20],2+ [80] V (Fig. 6) .</p><p>Bioacoustics. The description is based on calls recorded by Julian Dring at Camp 3, Gunung Mulu National Park, Sarawak, on 8 September 1977 at 1700 h. Both L. gunungensis sp. nov. and L. nephophilus sp. nov. have been collected at Camp 3 by Julian Dring. The two species differ markedly in the extent of webbing (Fig. 6, Table 4). In an unpublished list provided together with recordings of the calls of some Southeast Asian frogs, J. Dring noted that the call was recorded from the “short-webbed form” of L. palavanensis, which we describe as L. nephophilus sp. nov. He did not collect a voucher specimen for the call, but stated that BMNH 1978.213 had the same call type. This specimen, a female, however, has extensive webbing and was referred by us to L. gunungensis sp. nov. We therefore tentatively assign the recorded call to the short-webbed L. nephophilus sp. nov. but point out that this assignment needs to be confirmed by future field work. The advertisement calls of males (n=3) are sharp chirps, 75–96 ms in duration, at a dominant frequency of 2.5–3.0 kHz, and with a marked frequency modulation (Fig. 11D; Table 3). The calls are given in bouts of up to 21 calls, in which calls are separated from each other by an interval of 135–383 ms. A sound spectrogram and a waveform obtained from the same recording were depicted in Malkmus et al. (2002: 147; as L. palavanensis, the population herein described as L. kinabaluensis sp. nov., see above), erroneously stating that it had been recorded at an elevation of 150 m in Gunung Mulu National Park.</p><p>Distribution. The species has been recorded from montane rainforest at elevations between 1,200 and at least 2,100 m on Gunung Mulu and Gunung Murud in eastern Sarawak (Figs. 8, 9). The species occurs sympatrically with L. gunungensis sp. nov. in the area around Camp 3 on Gunung Mulu, Sarawak.</p><p>Ecology. We tentatively assign a tadpole (BMNH 1978.216) to this species, which according to the catalog entry had been collected “in a small silty pool, between rocks on a dry stream bed” at Camp 3 in Gunung Mulu National Park. The tadpole was included in the description of the tadpole of “ Rana palavanensis ” (sensu lato) by Inger (1985). Free-swimming tadpoles of this species in advanced stages were collected at Camp 3 (lot field number 367; A. Haas, pers. comm.) and Camp 4 (NMBE 1065176–1065177), Gunung Mulu National Park. Juveniles were observed in leaf litter away from any water body on the trail along the mountain ridge between Camps 3 and 4. The number and size of the ripe ova of the females indicate small clutch sizes. Other details of the reproduction of the species are unknown, but we expect it to reproduce in a similar way to the closely related species for which the reproduction mode is known.</p><p>Etymology. The species epithet is composed of the Ancient Greek noun νέφος [néphos], meaning cloud, and the Latinized suffix -philus (from Ancient Greek φιλεῖν [phileĩn], meaning “to love”); meaning “the one who loves clouds”, in allusion to the habitat of the new species in montane cloud forests of Gunung Mulu and Gunung Murud.</p><p>Suggested English name. Cloud-Forest Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFF0FFD4FF34FC09FCCAF87A	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFFAFFD3FF34FF50FCE2F87C.text	62199F50FFFAFFD3FF34FF50FCE2F87C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes oreibates Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes oreibates sp. nov.</p><p>Rana palavanensis (nec Rana palavanensis Boulenger, 1894)— Boulenger 1920: 59 (partim); van Kampen 1923: 182 (partim); Smith 1925: 32.</p><p>Rana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1966: 222 (partim).</p><p>Limnonectes palavanensis (nec Rana palavanensis Boulenger, 1894)— Zainudin et al. 2017: 106.</p><p>Holotype. NMBE 1061519, adult female, from Borneo Highlands Resort (1.116667°, 110.216667°; 990 m), Gunung Penrissen, Sarawak, Malaysia, collected on 10 March 2012 by S. T. Hertwig, A. Jankowski, Y. M. Pui (Fig. 3E).</p><p>Paratypes. BMNH 1928.11.4, adult female, from Mt. Penrissen, “ N. Borneo ” [= Sarawak], Malaysia, “pres. by M. Smith” (probably collected by Eric Mjöberg; Smith 1925) . UNIMAS P1186, adult male, collected on 10 September 2015 by Y. M. Pui; UNIMAS P1193, juvenile, collected on 14 September 2015 by Y. M. Pui and I. Das, both from Summit Trail to Gunung Penrissen, below Batu Panggah, Sarawak, East Malaysia (Borneo) . UNIMAS P1192, adult male, from an unnamed forest trail within Borneo Highlands Resort area, Gunung Penrissen, Sarawak, East Malaysia (Borneo), collected on 14 September 2015 by Y. M. Pui and I. Das (Fig. 7E) .</p><p>Referred specimen. UNIMAS 9737, adult male, from the edge of an intermittent stream at Batu Panggah, Borneo Highlands Resort, collected by I. Das on 27 August 2021 .</p><p>Diagnosis. Limnonectes oreibates sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing reduced; supratympanic fold weakly curved; pineal spot distinctly discernible, anterior to level of anterior upper-eyelid edge; vomer ridges short, widely separated from each other; postpalpebral ridge weakly developed; lumbar and median ridges absent; size relatively large with SVL of adult males 32.5–33.2 mm, of adult females 35.9–37.1 mm. The species differs from all other species of the group by at least 5.8% in the sequence of the 16S barcoding gene. The advertisement call is a slow chirp, 174–317 ms in duration with a dominant frequency of 1.4–1.6 kHz with weak frequency modulation, given in call bouts of up to 11 calls with an intercall interval of 381–775 ms. The species occurs in montane rainforest at around 1,000 m.</p><p>Description of holotype. Adult female; SVL 35.9 mm; body sturdy, widest at temporal region, slightly tapering to groin (Fig. 3E); head large (HL/SVL 0.38, HW/SVL 0.39), slightly wider than long (HW/HL 1.04); snout moderately long (SL/HL 0.45), rounded in both dorsal view and lateral profile, wider than long (SL/EE 0.94); canthus rostralis distinct between eye and nostril, weakly concave between eye and nostril in dorsal view; loreal region oblique; nostrils oval, directed laterally; situated much closer to tip of snout than to eye (EN/NS 1.21), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.33); eye directed anterolaterally, protruding, moderately large (ED/HL 0.32), its diameter much shorter than snout (ED/SL 0.71); interorbital distance greater than upper eyelid width (IO/EW 1.17) and smaller than internarial distance (IO/NN 0.74); pineal spot distinctly discernible, anterior to level of anterior upper-eyelid edge; tympanum and its annulus distinctly visible; tympanum separated from eye by little more than half its diameter (ET/TD 0.57); tympanum diameter smaller than eye diameter (TD/ED 0.55); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, their lateral half covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, short, separated from choana by two-thirds length of individual process, from each other by distance subequal to length of individual process (Fig. 4E); tongue moderately long and narrow, bilobed for about one-seventh of its length, free distally for about two-thirds its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs finely shagreened; lateral surfaces of trunk finely areolate with sparsely scattered small tubercles; dorsolateral ridge prominent, from posterior end of upper eyelid to groin; postpalpebral ridge weakly developed; lumbar and median ridges absent; horseshoe-shaped dermal ridge low, in middle of back just behind level of arms; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, weakly curved (Fig. 5E); few enlarged tubercles on lateral edge of upper eyelid; ventral side of head smooth; transverse chest fold between arm insertions distinct; ventral side of trunk weakly areolate, almost smooth; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: II &lt;IV &lt;I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, flat, about two-fifths length of metacarpal of Finger I; inner and outer palmar tubercles fused proximally, bipartite for about one-fifth of its length on distal end, otherwise rounded, distinct, covering proximal one-third of metacarpals of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.80); heel reaching beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.57), slightly longer than thigh (TFL/THL 1.06); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.88); relative length of toes: I &lt;II &lt;III &lt;V &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1.5/2+ II 1+/2.5 III 1.5/3 IV 3/1.5 V (Fig. 6); narrow dermal flap on preaxial side of Toe I between metatarsal tubercle and disk and on postaxial side of Toe V, widest from proximal end of metacarpus to proximal subarticular tubercle, continues to disk; inner metatarsal tubercle prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life (based on paratype UNIMAS P1186; Fig. 7E) Dorsum mid-brown, with dark interorbital bar demarcating brownish-pink snout from darker dorsum; broad (ca. 2.0 mm) yellowish-cream stripe extending from snout-tip to above cloaca; middorsal stripe edged with black laterally, especially around snout region and at posterior portion terminating with two laterally placed black blotches; dorsolateral fold edged laterally with narrow black stripe; flanks pale grey with faint, scattered mottling; supratympanic fold grey-black; tympanum pinkish yellow; supra- and infralabials blackish-grey, with narrow cream-coloured bars, sometimes incomplete, forming blotches; axillary region, from area behind tympanum to about level of scapula brown-grey, distinctly darker than rest of flanks; upper surfaces of fore- and hindlimbs brownish-pink, with narrow dark crossbands restricted to lower arm and on thigh and shank; digits yellowish-pink with dark crossbars; venter pale, generally unpatterned, except edges of infralabial region, fore- and lower hindlimbs, and on manus and pes, these regions densely pigmented with grey; yellow base colour of lower limbs intense; rest of venter such as gular and abdominal region cream coloured; pupil black; dark brown Maltese cross-like pattern on iris, centred around pupil; iris yellowish-brown, paler peripherally.</p><p>In preserved state (based on holotype; Fig. 3E), dorsum light brown in centre, lightening to brownish light grey towards dorsolateral folds; interorbital bar wide, blackish brown; supratympanic fold black; narrow black line along dorsolateral fold; dark brown stripe on flanks along dorsolateral fold, anteriorly wider, dark brown and occupying whole area between supratympanic and dorsolateral folds, posteriorly lighter brown and narrower, with irregular ventral edge throughout; flanks with bluish tinge; horseshoe-shaped tubercle dark brown; upper eyelids light blue; tympanum and area around yellowish brown; sides of snout mid-brown; dorsal part of snout and limbs yellowish light brown; lower arms, fingers, legs and toes with regular dark brown narrow crossbars; toe webbing dark brown; ventral side of head, chest, arms and legs yellowish cream-white; faint, small dark brown spots in infralabial region; throat with faint greyish mottling; abdomen hardly pigmented, blueish white; postaxial edge of lower arms, palmar side of hand and plantar side of feet including webbing dark brown; postaxial side of thighs and shanks with dark brown mottling.</p><p>Variation. The paratypes match the holotype in general appearance and proportions. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. SVL of adult males is 32.5–33.2 mm, of adult females 35.9–37.1 mm (Table 3). Pedal webbing variation is I 1.5/2+ II 1+/2.5 III 1.5/3 IV 3/1.5 V (Fig. 6).</p><p>Bioacoustics. Advertisement calls of males (n=2: UNIMAS P1186, P1192) are slow chirps, 174–317 ms in duration with a dominant frequency of 1.4–1.6 kHz, and with weak frequency modulation (Fig. 11; Table 3). They are given in call bouts of up to 11 calls, in which individual calls are separated from each other by intervals of 381–775 ms (Fig. 11E; Table 3).</p><p>Distribution. So far, the species is known only from Gunung Penrissen in western Sarawak (Fig. 8), where it has been found at elevations between about 900 and 1,100 m (Fig. 9).</p><p>Ecology. The number and size of the ripe ova of the females indicate small clutch sizes. A male (UNIMAS 9737) was found carrying tadpoles at the edge of an intermittent stream at Batu Panggah on 27 August 2021.</p><p>Etymology. The species epithet derives from the Ancient Greek noun ορειβάτης [oreibátes], meaning “mountaineer”; in allusion to the type locality of the new species on top of Gunung Penrissen.</p><p>Suggested English name. Penrissen Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFFAFFD3FF34FF50FCE2F87C	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFF8FFECFF34FF50FCF7F9A0.text	62199F50FFF8FFECFF34FF50FCF7F9A0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes phylax Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes phylax sp. nov.</p><p>Rana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894)— Inger 1966: 222 (partim). Rana palavanensis (nec Rana palavanensis Boulenger, 1894)— Inger &amp; Voris 1988: 1060; Das 1995: 161. Limnonectes palavanensis (nec Rana palavanensis Boulenger, 1894)— Das 2007: 53; Grafe &amp; Keller 2009: 28; Grafe et al.</p><p>2010: 55; Goyes Vallejos &amp; Ahmad Sah 2017: 608; Inger et al. 2017: 87 (partim); Goyes Vallejos et al. 2017: 1, 2018:</p><p>2473, 2019: 29.</p><p>Holotype. NMBE 1072362, adult female, from Rhino Pool, Danum Valley Conservation Area, Sabah, Malaysia, collected on 2 March 2015 by S. T. Hertwig et al. (Fig. 3F).</p><p>Paratype. NMBE 1073856, adult female, from Loop Walk (4.3742°, 115.43844°; 475 m), Merarap Hot Spring Lodge, Sarawak, Malaysia, collected on 1 March 2016 by J. Flury, A. Haas, S . T. Hertwig, Pui Y. M. and N. Reichen .</p><p>Referred material. NMBE 1064757, tadpole, from Gunung Mulu National Park (100 m), Sarawak, Malaysia, collected on 11 March 2006 by I. Das,A. Haas and S . T. Hertwig . NMBE 1065198, tadpole, from Tawau Hills National Park, Sabah, Malaysia, collected on 13 August 2007 by I. Das, C. Garratt, A. Haas and K. Hee . NMBE 1064089, tadpoles, from Sungai Tawau (4.461083°, 117.920764°; 828 m), Tawau Hills National Park, Sabah, Malaysia, collected on 10 August 2007 . NMBE 1072710, tadpoles, from Tembaling Waterfall (4.941332°, 117.807974°; 265 m), Danum Valley Conservation Area, Sabah, Malaysia, collected on 3 March 2015 by J. Flury, A. Haas and S . T. Hertwig . UBDM-GG010, adult female; UBDM-GG011, GG011, adult male; both from Ashton Trail, towards Sungai Mata Ikan, near Kuala Belalong Field Studies Centre, Ulu Temburong National Park, Temburong, District, Brunei Darussalam, collected on 23 July 2012 by J. Goyes Vallejos. UBDM-GG012, adult male, collected on 16 October 2014; UBDM-GG013 –GG014, two adult males, UBDM-GG015–017, three adult females, all collected on 19 October and 19 November 2014; all from near Kuala Belalong Field Studies Centre, Ulu Temburong National Park, Temburong, District, Brunei Darussalam, collected by T. U. Grafe and J. Goyes Vallejos. SP 894, adult male, from Purulon Camp, Tenom District, Crocker Range, Sabah, Malaysia, collected on 23 June 1989 by “ R. F. Inger et al.”. SP 895, adult male, from “Mondolong” [= Mendolong] , SFI, Sipitang, Sabah, Malaysia, collected on 2 August 1989 by “ R. F. Inger et al.”. SP 2242 (tag on specimen) or 2243 (jar label), adult male, from Long Pasia (4.40°, 115.72°; 1,000 m), Sipitang, Sabah, Malaysia, collected on 6 May 1998 by “Fred, Paul, et al.”. SP 2649, adult male, from Tawau Hills Park, Sabah, Malaysia, collected on 23 May 2000 by “Maklarin [Lakim], Alim [ Biun] &amp; Benedik ” .</p><p>Diagnosis. Limnonectes phylax sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing moderately extensive; supratympanic fold strongly curved; pineal spot hardly discernible; vomer ridges long, narrowly separated from each other; postpalpebral ridge well developed, from upper eyelid to anterior level of pelvis; median ridge low from tip of snout to horseshoe-shaped ridge, much more conspicuous from horseshoe-shaped ridge to vent; lumbar ridge well developed; size relatively small with SVL of adult males 26.0– 28.5 mm, of adult females 30.1–34.5 mm. The species differs from all other species of the group by at least 6.6% in the sequence of the 16S barcoding gene. The advertisement call is a long rising trill, consisting of 21–39 notes, lasting 1.4– 2.2 s, at a dominant frequency of 1.4–2.1 kHz with a weak frequency modulation and an amplitude maximum in the first third of the call. The species occurs in lowland rainforest up to 830 m elevation.</p><p>Description of holotype. Adult female; SVL 32.0 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3F); head large (HL/SVL 0.35, HW/SVL 0.35), about as long as wide (HW/HL 1.01); snout long (SL/HL 0.47), subacuminate in dorsal view, rounded and slightly protruding in lateral profile, slightly wider than long (SL/EE 0.90); canthus rostralis moderately distinct between eye and nostril, almost straight-lined from eye to nostril in dorsal view; loreal region oblique; nostrils rounded, directed dorsolaterally; situated slightly closer to tip of snout and to eye (EN/NS 1.05), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.39); eye directed anterolaterally, moderately protruding, very large (ED/HL 0.34), its diameter shorter than snout (ED/SL 0.72); interorbital distance about as wide as upper eyelid (IO/EW 1.06) and smaller than internarial distance (IO/NN 0.74); pineal spot hardly discernible, covered by skin, as small, low elevation posterior to level of anterior upper-eyelid edge; tympanum and its annulus visible, its posterodorsal edge in contact with supratympanic fold, its posteroventral edge covered by thick skin; tympanum separated from eye by two-fifth of its diameter (ET/TD 0.40); tympanum diameter smaller than eye diameter (TD/ED 0.62); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located far anterolaterally at margins of roof of mouth, lateral third covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, long, separated from choana by about half length of individual process, from each other by less than width of individual process, almost in contact medially (Fig. 4F); tongue short and broad, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs and lateral surfaces of trunk shagreened; enlarged tubercles sparsely scattered on upper eyelid and on dorsum; low, horseshoe-shaped dermal ridge in middle of back just behind level of arms; seven longitudinal dermal ridges running more or less parallelly to each other along back; dorsolateral ridge most prominent, from posterior end of upper eyelid to groin; postpalpebral ridge well developed, from upper eyelid to anterior level of pelvis; median ridge quite indistinct and very low from tip of snout to horseshoe-shaped ridge, continuing much more conspicuous from horseshoe-shaped ridge to vent; lumbar ridge well developed, from horseshoe ridge to vent, interrupted in posteriormost part; supratympanic fold thick and conspicuous, extending from posterior end of orbit, strongly curved, initially directed posterodorsally then turning sharply to arm insertion, straight-lined in its posterior fourth of length (Fig. 5F); ventral side of head, trunk and limbs smooth; transverse chest fold between arm insertions distinctly expressed; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.25); tips of fingers rounded, distinctly enlarged into disks; relative length of fingers: II = IV &lt;I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle oval, small, about two-fifth length of metacarpal of Finger I; inner and outer palmar tubercles fused proximally, forming roughly U-shaped, low tubercle on proximal third of metacarpals of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.82); heel reaching distinctly beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.58), slightly longer than thigh (TFL/THL 1.06); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.91); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1+/2 II 1+/2+ III 1+/2.5 IV 3-/1.25 V (Fig. 6); narrow dermal ridge on preaxial side of Toe I and on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life (Fig. 7F), dorsum light brown; horseshoe-shaped tubercle anteriorly coloured like dorsum with blackish brown posterior edge; dorsolateral fold with very narrow reddish line along medial edge and blackish brown lateral edge, black edge wider in posterior part; interorbital bar blackish brown with narrow cream anterior edge, lightening towards posterior edge to mid-brown, continued onto posterior part of upper eyelids; dorsal face of head anterior to interorbital bar as well as sides of snout greyish light brown; tympanic area bright yellowish, mottled with light brown; supratympanic fold and area above tympanum dark grey; flanks yellowish in anterior part, light grey in posterior part, with two dark brown spots in the middle; few small white dots at side of head, tympanic area and anterior part of flanks; arms pale yellow with few dark brown irregularly shaped spots; fingers dark brown with yellow spots; legs reddish light brown with greyish light brown crossbars; crossbars narrower and more frequent on thigh than on crus; ventral side of head, chest, arms, and legs bright yellow; abdomen pale yellow; dark brown speckling along postaxial side of lower arm and tarsus, and on preaxial and postaxial sides of thigh and crus; palmar side of hand and plantar side of foot dark brown.</p><p>In preserved state, dorsum of holotype grey overall with light brown tinge in anterior half of trunk and on crus. Reddish lines faded to brown; yellow colouration on arm faded to brownish grey; greyish brown crossbars on legs changed to light brown; upper eyelids with bluish tinge (Fig. 3F); ventral side of head, chest, and legs pale yellow; abdomen off white with bluish tinge in centre, pale orange on flanks; palmar side of hand and plantar side of feet light brown.</p><p>Variation. The paratypes match the holotype in general appearance. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. SVL of adult males is 26.0– 28.5 mm, of adult females 30.1–34.5 mm (Table 2). Inger &amp; Voris (1988) recorded 25.2–28.2 mm SVL for males and 26.3–32.3 mm for females from Danum Valley.Pedal webbing variation is I 1 [63],1+ [25],1.25 [13] /2 II 1 [50],1+ [50] /2 [37],2+ [37],2.25 [13],2.5 [13] III 1 + [25],1.25 [37],1.5 [37] /2.5 [3 7],3- [63] IV2.75 [13],3- [87] /1 [37],1+ [25],1.25 [37] V (Fig. 6) .</p><p>Bioacoustics. We have analyzed calls recorded at Camp 5, Gunung Mulu National Park (Sarawak, not collected), Ulu Temburong National Park (Brunei, UBDM-GG011–14), Sepilok Forest Reserve ( Sabah, not collected) and Tawau Hills National Park ( Sabah, not collected). The call characteristics did not differ between the sites. The advertisement call of males (n=7) is a long rising trill, containing 21–39 notes and lasting 1.4– 2.2 s. Individual notes last 36–45 ms and are repeated at a rate of 15 notes/s. Dominant frequency is at 1.4–2.1 kHz, with a weak upward modulation from the beginning to the end or the call (Fig. 11B; Table 3). The amplitude initially increases rapidly, reaches its maximum within the first third of the call and subsequently slowly and weakly decreases until the end of the call (Fig. 11B). Females often call in response to the advertisement call of the male with very soft single-note squeaks or chirps, lasting on average 56.8 ± 9.0 ms, with a dominant frequency of 1,608 ± 67 Hz (Goyes Vallejos et al. 2017). Males use softer and shorter single-note courtship calls with an average duration of 309 ± 126 ms and an average dominant frequency of 1,752 ± 86 Hz in response to female calls (Goyes Vallejos et al. 2017). The call description as “a short, high-pitched trill” by Inger &amp; Stuebing (1989) for specimens from Sabah roughly matches the call of L. phylax sp. nov ..</p><p>Distribution. So far, the occurrence of species has been confirmed for a few locations between 100 and 830 m in eastern Sarawak (Gunung Mulu National Park, Merarap), Brunei Darussalam (Ulu Temburong National Park), and Sabah (Danum Valley, Tawau Hills National Park, Sepilok Forest Reserve) (Fig. 8). We tentatively refer specimens from further locations in Sabah (Purulon, Sipitang) to the species. We expect the species to be widely distributed in lowland rainforests of northern Sarawak, Brunei, Sabah, as well as northeastern Kalimantan. The species occurs sympatrically with L. finchi in Sabah, and parapatrically with L. sarawakensis sp. nov. in the lowlands of Gunung Mulu National Park, Sarawak, where the species seem to be distributed in different forest types ( L. phylax sp. nov. in dipterocarp forest and karst forest, L. sarawakensis sp. nov. in alluvial forest); and with L. kinabaluensis sp. nov. in Tawau Hills National Park, Sabah, where the two species differ in elevational distribution with the latter occupying higher elevations (Fig. 9).</p><p>Ecology. Eggs are deposited in moist leaf litter on top of dead leaves and guarded by the male (Inger &amp; Voris 1988). Clutches consist of 5– 21 eggs and tadpoles hatch at Gosner stage 24 or 25 9–11 days after oviposition (Inger &amp; Voris 1988; Goyes Vallejos et al. 2018). Males do not call during this time, nor do they care for more than one clutch at a time (Goyes Vallejos et al. 2018). After hatching, the tadpoles crawl onto the male’s back and are transported to water bodies (Inger &amp; Voris 1988, Inger et al. 2017, Goyes Vallejos et al. 2018; Fig. 11). Observations of tadpole transport in the field indicate that all tadpoles from a clutch are retrieved at once and that the male stimulates synchronous hatching (Goyes Vallejos et al. 2018). Adults of the species have been collected in the same forests blocks as L. finchi in Danum Valley (Inger &amp; Voris 1988). The species is preyed on by the Rough-backed Snake ( Xenodermus javanicus) (Goyes Vallejos &amp; Ahmad Sah 2017).</p><p>Etymology. The species epithet derives from the Ancient Greek noun φύλαξ [phylax], meaning “guard” or “guardian”, and is used as a noun in apposition; in allusion to the egg-guarding behaviour of the males of the species.</p><p>Suggested English name. Smooth Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFF8FFECFF34FF50FCF7F9A0	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFC6FFE9FF34F9B8FCFDFE5C.text	62199F50FFC6FFE9FF34F9B8FCFDFE5C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes sarawakensis Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes sarawakensis sp. nov.</p><p>Rana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1966: 222 (partim); Inger et al. 1986: 523. Rana palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1985: 52 (partim); Inger &amp; Voris 1988: 1060 (partim). Limnonectes palavanensis (nec Rana palavanensis Boulenger, 1894)— Das et al. 2016: 19 (partim); Dehling &amp; Dehling 2021:</p><p>Table A2.</p><p>Holotype. NMBE 1057455, adult female, from Lambir Hills National Park (4.2°, 114.0°; 100–400 m), Sarawak, Malaysia, collected in September 2010 by A. Haas, S. T. Hertwig and A. Jankowski (Fig. 3G).</p><p>Paratypes. NMBE 1057451, adult male, collected on 8 September 2010 by A. Haas, S . T. Hertwig, A. Jankowski, NMBE 1069412, adult male, collected on 14 March 2014 by S . T. Hertwig et al., both from Lambir Hills National Park, Sarawak, Malaysia . NMBE 1101233–1101234, two adult males, NMBE 1101235, juvenile; all from Camp Koyan (3.342366°, 114.177573°; 822 m), High Plateau, Gunung Dulit, Sarawak, Malaysia , collected on 8 February 2019 by L. Etter, S . T. Hertwig and F. Heussler . NMBE 1101236, adult male, from Summit Trail (2.655489, 112.903817; 185 m), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=112.90382&amp;materialsCitation.latitude=2.655489" title="Search Plazi for locations around (long 112.90382/lat 2.655489)">Bukit Kana National Park</a>, Sarawak, Malaysia , collected on 15 March 2019 by L. Etter, S . T. Hertwig and F. Heussler . NMBE 1101217–1101219, three adults of undetermined sex, from unnamed stream at 537 m (3.096895°, 114.2673°; 537 m), Camp Balah, Gunung Dulit, Sarawak, Malaysia , collected on 24 February 2019 by L. Etter, L. Fehland, S. T. Hertwig and F. Heussler . NMBE 1101238–1101240, three adults of undetermined sex, from sidestreams and ponds near camp (1.960155°, 114.783815°; 495 m), Long Unai, Sarawak, Malaysia , collected on 13 March 2018 by S. Cilloni, L. Etter, L. Fehland, S. T. Hertwig, B. Minder and Y. M. Pui .</p><p>Referred specimens. NMBE 1065274, tadpole, from alluvial forest near headquarters, Gunung Mulu National Park, Sarawak, Malaysia, collected on 24 March 2009 by A. Haas . ZRC 12728, adult male, from Nanga Tekalit (1.63°, 113.58°), Sarawak, Malaysia, collected on 28 March 1984 by R. F. Inger and P. Walker .</p><p>Diagnosis. Limnonectes sarawakensis sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toe webbing extensive; supratympanic fold strongly curved; pineal spot weakly discernible, anterior to level of anterior upper-eyelid edge; vomer processes long, separated from choana and from each other by about half length of individual process; postpalpebral ridge well developed, from upper eyelid to anterior level of pelvis; median ridge indistinct from level of posterior upper-eyelid edge to horseshoe-shaped ridge, more conspicuous from horseshoe-shaped ridge to vent; lumbar ridge well developed, interrupted; size relatively small with SVL of adult males 25.2–29.7 mm, of adult females 30.1–34.4 mm. The species differs from all other species of the group by at least 5.8% in the sequence of the 16S barcoding gene. The advertisement call is a long rising trill, consisting of 26–32 notes, lasting 1.6– 2.0 s, at a dominant frequency of 1.4–1.8 kHz with a weak frequency modulation and an amplitude maximum at the end of the call. The species is distributed in lowland and submontane rainforests of central and northern Sarawak, between 60 and 1,000 m.</p><p>Description of holotype. Adult female; SVL 31.2 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3G); head large (HL/SVL 0.36, HW/SVL 0.37), about as long as wide (HW/HL 1.03); snout long (SL/HL 0.45), subacuminate in dorsal view, rounded and slightly protruding in lateral profile, slightly wider than long (SL/EE 0.92); canthus rostralis moderately distinct between eye and nostril, slightly concave from eye to nostril in dorsal view; loreal region oblique; nostrils rounded, directed dorsolaterally; situated closer to tip of snout and to eye (EN/NS 1.14), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.30); eye directed anterolaterally, moderately protruding, very large (ED/HL 0.35), its diameter shorter than snout (ED/SL 0.78); interorbital distance about as wide as upper eyelid (IO/EW 1.02) and smaller than internarial distance (IO/NN 0.73); pineal spot weakly discernible, anterior to level of anterior upper-eyelid edge; tympanum and its annulus visible, its posterodorsal edge in contact with supratympanic fold; tympanum separated from eye by one-third of its diameter (ET/TD 0.35); tympanum diameter smaller than eye diameter (TD/ED 0.59); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, subtriangular, located far anterolaterally at margins of roof of mouth, lateral third covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, long, separated from choana and from each other by about half length of individual process (Fig. 4G); tongue short and broad, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs and lateral surfaces of trunk rough; enlarged tubercles sparsely scattered on the upper eyelid and on the dorsum; low, horseshoe-shaped dermal ridge in middle of back just behind level of arms; seven longitudinal dermal ridges running more or less parallelly to each other along the back; dorsolateral ridge most prominent, from posterior end of upper eyelid to groin; postpalpebral ridge well developed, from upper eyelid to anterior level of pelvis; median ridge indistinct from level of posterior upper-eyelid edge to horseshoe-shaped ridge, continuing more conspicuous from horseshoe-shaped ridge to vent, interruptedly above pelvis; lumbar ridge well developed, running interrupted from horseshoe ridge to vent; supratympanic fold thick and conspicuous, extending from posterior end of orbit, strongly curved, initially directed posterodorsally then turning sharply to arm insertion (Fig. 5G); ventral side of head smooth; transverse chest fold between arm insertions weakly distinct; ventral side of trunk smooth, evenly scattered with enlarged tubercles; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.23); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: II = IV &lt;I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle oval, small, about two-fifth length of metacarpal of Finger I; inner and outer palmar tubercles fused proximally, forming roughly U-shaped, low tubercle on proximal third of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.76); heel reaching slightly beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.56), slightly longer than thigh (TFL/THL 1.09); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot shorter than tibiofibula (FOT/TFL 0.88); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1/2 II 1/2+ III 1+/2.25 IV 2.5/1+ V (Fig. 6); narrow dermal ridge on preaxial side of Toe I and on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, about half length of metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life (Fig. 7G), dorsum light brown; horseshoe-shaped tubercle coloured like dorsum with blackish brown posterior edge in centre and small spot at lateral ends; dorsolateral fold with very narrow creamish line along medial edge and blackish brown lateral edge, black edge wider in posterior part; interorbital bar black with narrow light grey anterior edge, continued onto posterior part of upper eyelids; dorsal face of head anterior to interorbital bar greyish light brown with black speckling; sides of head and tympanic area light brown with large black supralabial flecks and few small white dots; supratympanic fold black with white dots; area above tympanum and anterior part of flanks light brown with small and larger dark spots; posterior part of flanks greyish brown with off-white mottling; arms orange brown with few dark brown irregularly shaped spots; fingers I and II bright yellow, fingers II and IV pale yellow, all with dark brown spots; legs reddish light brown with greyish light brown crossbars; crossbars narrower and more frequent on thigh than on crus; ventral side of head, arms, and legs dark yellow; dark brown spots along mandibular edge; chest and abdomen bright yellow; greyish brown speckling along postaxial side of lower arm and tarsus, and on preaxial and postaxial sides of thigh and crus; palmar side of hand and plantar side of foot dark brown.</p><p>In preserved state, dorsum of holotype brownish grey overall with more distinct light brown colour along dorsal dermal ridges and on postaxial half of thigh Black dorsal pattern elements lightened to dark brown; orange colouration on arm, yellow colouration on hands, and reddish colouration on legs faded to brownish grey; crossbars on legs changed to mid-brown; upper eyelids with bluish tinge (Fig. 3G); ventral side of head, chest, arms and legs pale reddish cream; spots along mandibular edge light brown; abdomen hardly pigmented, mottled bluish and yellowish; palmar side of hand and plantar side of feet light brown.</p><p>Variation. The paratypes match the holotype in general appearance. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. SVL of adult males is 25.2–29.7 mm, of the adult female 31.2 mm (Table 2). Inger &amp; Voris (1988) recorded 26.8–31.6 mm SVL for males and 26.1–34.4 mm for females from Nanga Tekalit. Pedal webbing variation is I 1 [71],1+ [29] /2 [57],2+ [43] II 1 [71],1+ [29] /2 [43],2+ III 1 +/2 [14],2+ [29],2.25 [14],2.5 [14],2.75 [29] IV 2 [14],2+ [14],2.5 [29],2.75 [14],3- [29] /1+ V (Fig. 6) .</p><p>Bioacoustics. The advertisement call of males (n=1) is a long rising trill, consisting of 26–32 notes and lasting 1.6– 2.0 s. Individual notes last 33–47 ms and are repeated at a rate of 15 notes/s (Table 3). Frequency is weakly modulated and continuously increases from the beginning to the end of the call, with the dominant frequency constantly rising from 1.4 to 1.8 Hz (Fig. 11C). The amplitude increases continuously from the beginning, reaching its maximum at the end of the call (Fig. 11C).</p><p>Distribution. The species is distributed in lowland and submontane rainforest in central and eastern Sarawak (Lambir Hills National Park, Gunung Mulu National Park, Gunung Dulit National Park, Bukit Kana National Park, Long Unai, Nanga Tekalit; Fig. 8), between 50 and 1,000 m (Fig. 9). The species occurs sympatrically with L. gunungensis sp. nov. on the High Plateau on Gunung Dulit, Sarawak, and parapatrically with L. phylax sp. nov. in the lowland forests of Gunung Mulu National Park, Sarawak (Fig. 9).</p><p>Ecology. Males of the species are known to transport up to about 30 tadpoles on their backs to water bodies (Inger et al. 1986). Tadpoles were found in pools of small intermittent streams, in low shallow buttress tank, in a water-filled hole in a log, in small isolated rain pools on forest floor (Inger et al. 1986, Inger &amp; Voris 1988) and in a shallow puddle with silky bottom and scattered leaves together with tadpoles of Occidozyga in alluvial forest in Gunung Mulu National Park (A. Haas, pers. comm.). Total clutch size of females was estimated at 35–50 (Inger et al. 1986), later corrected to half this number (Inger &amp; Voris 1988). Tadpole of the species were described by Inger (1966, 1985). Like in other species of the group, we expect the species to deposit eggs in moist leaf litter which are subsequently guarded by the males until the hatched tadpoles are transported to water bodies.</p><p>Etymology. The species epithet is composed of the name of the Malaysian State of Sarawak and the Latin suffix -ensis (“from”); meaning “the one from Sarawak ”, in allusion to the distribution of the new species.</p><p>Suggested English name. Sarawak Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFC6FFE9FF34F9B8FCFDFE5C	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFC3FFEBFF34FDE4FCF3FC60.text	62199F50FFC3FFEBFF34FDE4FCF3FC60.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes suboliferus Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes suboliferus sp. nov.</p><p>Rana palavanensis (nec Rana palavanensis Boulenger, 1894)— Boulenger 1920: 59 (partim); van Kampen 1923: 182 (partim). Rana microdisca palavanensis (nec Rana palavanensis Boulenger, 1894): Inger 1966: 222 (partim).</p><p>Limnonectes palavanensis (nec Rana palavanensis Boulenger, 1894)— Das et al. 2008: 152; Inger et al. 2017: 87 (partim).</p><p>Holotype. NMBE 1059945, adult female, from the “Frog Pond” (1.606397°, 110.188612°; 295 m), Kubah National Park, Matang Range, Sarawak, East Malaysia (Borneo), collected on 24 September 2009 by J. M. Dehling and D. M. Dehling (Fig. 3H).</p><p>Paratypes. NMBE 1056615, adult male from the type locality, collected on 9 April 2009 by J. M. Dehling . NMBE 1059943, 1059944, 1059946, 1059947, four adult males; same collection details as holotype . NMBE 1065317–1065318, two adults of undetermined sex, from Kampung Sebako (1.736725°, 109.737347; 20 m), at the foot of Gunung Pueh, Sarawak, Malaysia, collected on 16 August 2011 by A. Haas, Y. M. Pui and R. Sujang . NMBE 1057078, from stream behind frog pond (326 m), Kubah National Park, Sarawak, Malaysia, collected on 17 August 2010 by A. Haas and S . T. Hertwig . NMBE 1063852, from Matang, Sarawak, Malaysia, collected on 9 March 2005 by I. Das, N. Yaakob and J. Sukumaran . NMBE 1063936 from Matang Wildlife Centre (1.6094°, 110.1601°; 15 m), Kubah National Park, Sarawak, Malaysia, collected on 4 August 2006 by I. Das, G . V. A. Gee and A. Haas . NMBE 1064798, tadpole, from Kubah National Park, Sarawak, Malaysia, collected on 4 August 2006 by I. Das, G . V. A. Gee and A. Haas . NMBE 1064829, tadpole, from Kubah National Park (310 m), Sarawak, Malaysia, collected on 27 August 2006 by I. Das and A. Haas . NMBE 1064849, tadpole, from Kubah National Park (310 m), collected on 3 March 2007 by I. Das, A. Haas and T. Riehl . NMBE 1068207, NMBE 1068211, two adults of undetermined sex, from stream behind frog pond (326 m), Kubah National Park, collected on 10 August 2012 by A. Haas and J. Hofmann . NMBE 1068312, adult of undetermined sex from Kubah National Park (295 m), collected on 4 September 2009 by H. Dobbeck and F. Meyer . NMBE 1069798, tadpole, from Kubah National Park (310 m), collected on 7 September 2009 by I. Das, A. Haas and A. Jankowski . NMBE 1069822, tadpole, from Kubah National Park, collected on 17 August 2010 by A. Haas and S . T. Hertwig . NMBE 1101222, adult male, from Viewpoint Trail (1.695378°, 109.844108°; 144 m), collected on 17 March 2019 ; NMBE 1101223–1101225, three adult males, from Summit Trail (1.713522°, 109.836314°; 419 m), collected on 19 February 2019; all from Gunung Gading National Park, Sarawak, Malaysia, collected by L. Etter, S . T. Hertwig and F. Heussler .</p><p>Diagnosis. Limnonectes suboliferus sp. nov. belongs to the L. palavanensis species group, showing the typical characters of the group, including small size (SVL &lt;40 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum finely shagreened, with prominent longitudinal dorsolateral glandular ridges; interorbital distance equal to or larger than upper eyelid width; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from all other species of the group by a unique combination of morphological characters, including toes very extensively webbed; supratympanic fold curved; pineal spot usually well discernible, rarely indistinct, slightly in front of anterior upper-eyelid edge; vomer ridges long, widely separated from each other; postpalpebral ridge weakly developed, inconspicuous; lumbar and median ridges low and inconspicuous; ventral side of snout with evenly scattered, tiny, glandular pits in males; size relatively small with SVL of adult males 25.8–31.2 mm, of adult female holotype 35.5 mm. The species differs from all other species of the group by at least 6.2% in the sequence of the 16S barcoding gene. The advertisement call is a brief chirp, 132–231 ms in duration at a dominant frequency of 1.6–2.0 kHz with marked frequency modulation, given in bouts of up to 11 calls. The species occurs in lowland rainforest below 400 m.</p><p>Description of holotype. Adult female; SVL 35.5 mm; body moderately sturdy, widest at temporal region, slightly tapering to groin (Fig. 3H); head large (HL/SVL 0.39, HW/SVL 0.36), slightly longer than wide (HW/HL 0.93); snout moderately long (SL/HL 0.42), subacuminate in dorsal view, slightly protruding in lateral profile, slightly wider than long (SL/EE 0.94); canthus rostralis distinct between eye and nostril, almost straight-lined in dorsal view; loreal region oblique; nostrils oval, directed laterally; situated closer to tip of snout than to eye (EN/ NS 1.27), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.24); eye directed anterolaterally, moderately protruding, moderately large (ED/HL 0.31), its diameter shorter than snout (ED/ SL 0.75); interorbital distance greater than upper eyelid width (IO/EW 1.23) and smaller than internarial distance (IO/NN 0.76); pineal spot weakly discernible, slightly in front of level of anterior upper-eyelid edge; tympanum and its annulus distinctly and fully visible, not covered by supratympanic fold, its posterior edge concealed by thickened skin; tympanum separated from eye by one-third its diameter (ET/TD 0.33); tympanum diameter smaller than eye diameter (TD/ED 0.58); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located anterolaterally at margins of roof of mouth, not covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, long, separated from each other and from choana by distance about two-thirds of length of individual process (Fig. 4H); tongue moderately long and narrow, bilobed for about one-sixth of its length, free distally for about half its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs and lateral surfaces of trunk finely shagreened; small tubercles sparsely scattered medially of supratympanic fold and around upper eyelid; low horseshoe-shaped dermal ridge in middle of back just behind level of arms; dorsolateral ridge prominent, from posterior end of upper eyelid to groin; postpalpebral ridge weakly developed, inconspicuous, narrow and low from upper eyelid to just behind level of horseshoe-shaped ridge; lumbar ridge low, narrow and inconspicuous from horseshoe-shaped ridge almost to vent, interrupted above pelvis; median ridge inconspicuous, narrow and low from level of posterior upper-eyelid edge to vent; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, curved (Fig. 5H); ventral side of head smooth; indistinct transverse chest fold between arm insertions; ventral side of trunk weakly areolate; ventral side of limbs smooth; cloaca without dermal flap.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, slightly enlarged into disks; relative length of fingers: I = II = IV &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV larger and more prominent than distal ones; finger webbing absent; thenar tubercle distinct, oval, prominent, about one-third length of metacarpal of Finger I; inner and outer palmar tubercles fused proximally, forming large, roughly U-shaped, prominent tubercle on proximal half of Fingers II–IV.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.96); heel reaching about one SL beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.64), longer than thigh (TFL/THL 1.17); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot shorter than tibiofibula (FOT/TFL 0.87); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1-/1.75 II 1/2 III 1/2+ IV 2+/1 V (Fig. 6); narrow dermal ridge on preaxial side of Toe I and on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, about two-fifths length of metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life (Fig. 7H), dorsum mid-brown, darkening to dark brown between interorbital bar and horseshoe-shaped tubercle; horseshoe-shaped tubercle blackish brown with light brown fringe posteriorly; dorsolateral fold marked by narrow red line, bordered medially with narrow light brown band and laterally with wide black band; interorbital bar wide, black with narrow white anterior edge, and light brown posterior edge, continued onto posterior part of upper eyelids; dorsal face of head anterior to interorbital bar light brown with orange tinge and central black spot around orange pineal spot; sides of head dark brown with large black supralabial flecks; tympanic area yellow with small red dots anteriorly; supratympanic fold dark brown; flanks light brown with large black sports surrounded by light brown narrow fringe; light bluish dots speckled on ventral part of flanks; arms orange brown with small red dots and few large, black spots, widened to crossbars on lower arm and fingers; legs greyish mid-brown with blackish brown crossbars; crossbars more frequent on thigh and tarsus than on crus; toes proximally reddish brown, distally red; ventral side of head, arms, and legs dark yellow; dark brown spots along mandibular edge; chest and abdomen bright yellow; greyish brown speckling along postaxial side of lower arm and tarsus, and on preaxial and postaxial sides of thigh and crus; palmar side of hand and plantar side of foot dark brown.</p><p>In preserved state, dorsum of holotype light brown overall; dorsolateral ridge greyish light brown; horseshoe-shaped tubercle, interorbital bar and crossbands on arms and fingers dark brown; crossbars on legs mid-brown, almost faded on tarsus; upper eyelids with bluish tinge (Fig. 3H); ventral side of head, arms and legs pale yellow; spots on mandibular edge light brown; chest and abdomen off-white with bluish tinge; palmar side of hand and plantar side of feet light brown.</p><p>Variation. The paratypes match the holotype in general appearance and proportions. Males lack nuptial pads and have weak dermal folds parallel to posterior part of mandible and several evenly scattered, tiny, glandular pits on ventral side of head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. The pineal spot is well discernible in most paratypes. SVL of adult males is 25.8–31.2 mm (Table 2). Pedal webbing variation of paratypes is I 1- [73],1 [27] /2- [82],2 [18] II 1 [91],1+ [9] /2 III 1 [82],1+ [18] IV 2 [9],2+ [82],2.25 [9] /2 [9],2+ [55],2.25 [27], 2,5 [9] /1 [55],1+ [45] V (Fig. 6) .</p><p>Bioacoustics. Advertisement calls of males (n=3: NMBE 1059943, 1059946, and an uncollected male) are brief chirps, 132–231 ms in duration, emitted in bouts of up to 12 calls, in which individual calls are separated from each other by intervals of 311–903 ms (Fig. 11F; Table 3). The dominant frequency is at 1.6–2.0 kHz with a marked frequency modulation (Fig. 11F).</p><p>Distribution. The species is distributed in lowland rainforest in western Sarawak (Kubah National Park, Gunung Gading National Park, Gunung Pueh; Fig. 8) between 15 and 330 m (Fig. 9).</p><p>Ecology. We observed calling males and gravid females in distances of between 10 and 20 m from a permanent pond in secondary ridge-top kerangas forest in Kubah National Park. Adult males were also recorded calling, and observed guarding an egg clutch and carrying tadpoles (Fig. 10C&amp;D) in flat riverine forest in the same area.</p><p>Etymology. The species epithet is composed of the Latin noun suboles, meaning “offspring”, and the Latin suffix -ferus (from ferre = “to carry”), meaning “the one who carries”; and means “the one that carries the offspring”, in allusion to the males of the species carrying their tadpoles.</p><p>Suggested English name. Western Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFC3FFEBFF34FDE4FCF3FC60	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFC1FFE5FF34FB40FD03F834.text	62199F50FFC1FFE5FF34FB40FD03F834.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes finchi (Inger 1966)	<div><p>Limnonectes finchi (Inger, 1966)</p><p>Rana microdisca (nec Boettger, 1892) — Boulenger 1920: 57 (partim).</p><p>Rana microdisca finchi — Inger 1966: 222; Inger et al. 1986: 523.</p><p>Rana finchi — Inger &amp; Voris 1988: 1060; Inger &amp; Stuebing 1992: 46 (partim).</p><p>Limnonectes finchi — Inger et al. 2000: 16 (partim); Inger et al. 2017: 79 (partim).</p><p>Holotype. FMNH 77499, adult male, from Kalabakan, Tawau District, Sabah [Malaysia], collected 28 June 1956 by R. F. Inger.</p><p>Paratypes. FMNH 63297, 63350, 76570, 76575–76576, 76579–76581, 76583–76588, 76592–76593, 76597– 76599, 76601, 76603–76504, 76626, 77497, eight unsexed juveniles, two subadult males, eight adult males, and six adult females, FMNH 77564, 18 tadpoles, all collected by R. F. Inger at Deramakot in the Kinabatangan basin, at Kalabakan, and at the Sepilok Forest Reserve, North Borneo [Sabah, Malaysia] between 9 June 1950 and 28 June 1956. The paratype FMNH 76597 (adult male, RFI field # 1277 from “ Deramakot, Kinabatangan District, N [orth]- Borneo ”) has been exchanged with the Naturhistorisches Museum Wien, Austria, in 1962 and is deposited under NMW 16961 (Fig. 3I; Gemel et al. 2019) .</p><p>Referred specimens. NMBE 1072557, adult male, NMBE 1072558, adult female, from Tembaling Waterfall (4.941332°, 117.807974°; 265 m), Danum Valley Conservation Area, Sabah, Malaysia, collected in March 2015 by S. T. Hertwig et al .. NMBE 1072461, adult male, from Sulphur Hot Spring (4.4155°, 117.9012; 360 m), Tawau Hills NP, Sabah, Malaysia, collected 14 March 2015 by S. T. Hertwig et al .. NMBE 1072364, adult female, from Rhino Pool, Danum Valley Conservation Area, Sabah, Malaysia, collected 2 March 2015 by S. T. Hertwig et al .. NMBE 1072468–1072469, two adult females, from Galas Waterfall (4.41569°, 117.889795°; 397 m), Tawau Hills NP, Sabah, Malaysia, collected 12 March 2015 by S. T. Hertwig et al .. NMBE 1072719 and 1072724, of undetermined sex, both from Tembaling Waterfall (4.941332°, 117.807974°; 265 m), Danum Valley Conservation Area, Sabah, Malaysia, collected on 28 February and 1 March 2015 , respectively, by J. Flury, A. Haas and S . T. Hertwig . NMBE 1064091, of undetermined sex, from Sungai Tawau (4.461083°, 117.920764°; 828 m), Tawau Hills National Park, Sabah, Malaysia, collected on 11 August 2007 . NMBE 1072599, of undetermined sex, from Headquarters of Danum Valley Conservation Area (159 m), Sabah, Malaysia, collected on 7 March 2015 by J. Flury, A. Haas and S . T. Hertwig .</p><p>Diagnosis. Limnonectes finchi belongs to the L. finchi species group, showing the typical characters of the group, including moderate size (SVL of males to 39.0 mm, of females to 45.6 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum granular with interrupted, longitudinal dorsal rows of elongate tubercles; interorbital distance shorter than upper eyelid width; pineal spot distinctly visible; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species shows a unique combination of morphological characters that distinguish it from the other species in the group, including dorsum with moderately amount of more or less densely scattered enlarged tubercles, cloaca without tubercles; skin on ventral side of head and chest hardly thickened; chest fold indistinct; loreal region oblique; pineal spot oval, at level with anterior eye edge; vomer processes bearing 7–9 teeth, almost in contact to each other medially, separated from each other by less than one-sixth length of individual process and from choana by about one-fourth length of individual process; hindlimbs long, tibiotarsal joint reaching well beyond snout; first finger relatively short, about as long as second and fourth fingers); low dermal fold present on postaxial side of Toe V. The species differs from the other species of the group by at least 4.4% in the sequence of the 16S barcoding gene. The advertisement call is a long rising trill, consisting of 31–34 notes, lasting 2.0– 2.2 s, at a dominant frequency of 0.8–1.1 kHz with a weak frequency modulation and an amplitude maximum at the end of the call.</p><p>Description of adult male (NMBE 1072461). SVL 31.6 mm; body sturdy, widest at level of arm insertion, markedly tapering to groin (Fig. 3I); head large (HL/SVL 0.37, HW/SVL 0.37), as wide as long (HW/HL 1.00); snout moderately long (SL/HL 0.45), rounded in both dorsal view and profile, slightly projecting beyond lower jaw, as wide as long (SL/EE 1.01); canthus rostralis distinct between eye and nostril, straight-lined; loreal region oblique, moderately concave; nostrils rounded, directed dorsolaterally, situated closer to tip of snout than to eye (EN/NS 1.18), separated from each other by distance larger than distance between eye and nostril (NN/EN 1.27); eye directed anterolaterally, considerably protruding, large (ED/HL 0.35), its diameter shorter than snout (ED/SL 0.76); interorbital distance slightly smaller than upper eyelid width (IO/EW 0.96) and smaller than internarial distance (IO/NN 0.75); pineal spot distinctly visible, oval, at level of anterior eye edge; tympanum and its annulus distinctly visible; tympanum separated from eye by little more than one-third of its diameter (ET/TD 0.38); tympanum diameter smaller than eye diameter (TD/ED 0.68); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, subtrapezoid, located far anterolaterally at margins of roof of mouth, not covered by palatal shelf of maxilla in ventral view; vomer processes well developed, bearing teeth, large, almost in contact to each other medially, separated from each other by less than one-sixth length of individual process and from choana by about one-fourth length of individual process (Fig. 4I); tongue comparatively short, bilobed for about one-fifth of its length, free distally for about two-thirds its length; median lingual process absent; vocal sac apertures very small and inconspicuous, short, at corner of mouth.</p><p>Dorsal surfaces of head, trunk and limbs rough, with scattered enlarged tubercles; tubercles in anterior half of dorsum more or less longitudinally enlarged, forming indistinct, interrupted ridges; tubercles in posterior part of dorsum rounded; inverted-v-shaped dermal ridge in middle of back just behind level of arms; dorsal surfaces of forearm, hand, tarsus and foot shagreened; lateral surfaces of trunk areolate with very few scattered small tubercles; thick dorsal skin of trunk ending abruptly at level of posterior end of upper eyelid, forming indistinct transverse dermal edge between eyes; dorsal and lateral faces of head finely granular; supratympanic fold thick and conspicuous, extending from posterior end of orbit to insertion of arm, curved; few enlarged tubercles on posterior part of upper eyelid; very conspicuous and large tubercle just posterior to mouth opening; skin on ventral side of head smooth with weak dermal folds parallel to posterior part of mandible; skin smooth on chest and abdomen; skin on ventral side of head and chest hardly thickened compared to skin on abdomen; transverse chest fold between arm insertions indistinct; ventral side of limbs smooth; cloaca without dermal flap and without tubercles.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.26); tips of fingers rounded, swollen and enlarged into disks; disk on Finger IV slightly smaller than disk on other fingers; relative length of fingers: IV = II = I &lt;III; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV much larger and more prominent than distal ones; subarticular tubercle of Fingers I and II larger than proximal subarticular tubercles on Fingers III and IV; finger webbing absent; thenar tubercle distinct, oval, flat, about one-third length of metacarpal of Finger I; inner palmar tubercle small, rounded, flat, on proximal end of metacarpus of Fingers II and III; outer palmar tubercle about as long as inner palmar tubercle but more distinct, on proximal third of metacarpal of Finger IV, fused with inner palmar at proximal base, about half size of thenar tubercle; nuptial pads absent.</p><p>Hindlimbs sturdy, very long (LEG /SVL 1.86); heel reaching about one SL beyond tip of snout when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.60), longer than thigh (TFL/THL 1.11); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot slightly shorter than tibiofibula (FOT/TFL 0.91); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1+/2+ II 1+/2.25 III 1.25/3 IV 3+/1.25 V (Fig. 6); weak dermal fold on postaxial side of Toe V from proximal end of metacarpus to disk; inner metatarsal tubercle prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent.</p><p>Colouration. In life (Fig. 12), dorsum medium olive-greenish brown; larger tubercles on dorsum light to reddish brown; interorbital bar dark brown with narrow light brown anterior edge, continued onto posterior part of upper eyelids; dorsal face of head anterior to interorbital bar light olive-greenish brown with dark brown speckling; pineal spot whitish; sides of head to tympanic area with alternate bands of dark olive-greenish brown and greenish white bands; tympanum covered with dark brown spot; supratympanic fold reddish brown; flanks light greenish grey; arms brown-greenish grey with small light reddish brown tubercles; fingers light grey with dark grey blotches; legs greenish light brown with mid-brown crossbars and small, light reddish brown tubercles; crossbars narrower and more frequent on thigh than on crus; toes reddish crème-coloured; webbing dark olive green; ventral side of head, arms, and legs off-white; dark brown spots along mandibular edge; throat mottled dark brown. In preserved state, dorsum dark greyish brown; dorsal pattern elements dark brown (Fig. 3I); limbs including fingers and toes somewhat lighter brown; reddish life colouration faded to light brown; ventral side light grey.</p><p>Variation. The examined specimens match the paratype as well as the original description of the type series of L. finchi by Inger (1966). SVL of adult males is 31.6–39.7 mm, of adult females 36.3–45.8 mm (Table 2; see also Inger &amp; Voris 1988). Pedal webbing variation is I 1 [29],1+ [71] /1.75 [14],2 [57],2+ [29] II 1+/2 [14],2+ [86] III 1+ [29],1.25 [57],1.5 [14] /3- [57],3 [43] IV 3- [43],3 [43],3+ [14] /1+ [29],1.25 [28],1.5 [43] V (Fig. 6).</p><p>Bioacoustics. Two advertisement calls from a single uncollected male were recorded in Danum Valley, Sabah. The advertisement call is a long series of 31–34 notes with a total duration of 2.0– 2.2 s (Fig. 11A; Table 3). It is emitted after irregular intervals of several seconds. Note duration regularly increases from 31 ms in the first note of the call to 34 ms in the last note. Internote interval increases from 29 ms between the first two notes to 47 ms between the last two notes of a call. As a consequence, note repetition rate decreases from 16.6 notes/s between the first two notes to 12.4 notes/s between the last two notes of the call. Amplitude is strongly modulated and exhibits a regular increase from the beginning to the end of the call (Fig. 11A). Three prominent frequency bands are visible with prevalent bandwidth between 500 and 1500 Hz. There is a more or less regular frequency increase between individual notes from the beginning to the end of the call and another one within each note. In the third note of the call, the fundamental increases from 559 Hz to 602 Hz, the dominant frequency from 818 Hz to 861 Hz, and the third harmonic from 1,162 Hz to 1,248 Hz; in the last note of the call the fundamental increases from 559 Hz to 732 Hz, the dominant frequency from 861 Hz to 1,076 Hz, and the third harmonic from 1162 Hz to 1,421 Hz (Fig. 11A).</p><p>Distribution. Northeastern Borneo between sea level and around 1,100 m (Figs. 8, 9).</p><p>Ecology. The holotype was observed carrying tadpoles at the edge of a rain pool on a road through a logged area (Inger 1966). 18 tadpoles were collected and the total number of tadpoles was estimated to be twice that number (Inger 1966). Eggs are deposited in moist leaf litter on the underside of dead leaves and are guarded by the male (Inger et al. 1986, Inger &amp; Voris 1988). After hatching, the tadpoles crawl onto the male’s back and are then transported (Fig. 10G) to a pond or other small water body in which they develop until metamorphosis (Inger &amp; Voris 1988). Tadpoles are found in pools of rain water on the forest floor (Inger 1985). Tadpoles of the species were briefly described by Inger (1966) and in detail by Inger (1985).</p><p>Etymology. The species epithet is a patronym honouring O. C. Finch “for much assistance and pleasant companionship” to R. F. Inger in the field (Inger 1966).</p><p>Suggested English name. Rough Guardian Frog.</p></div>	https://treatment.plazi.org/id/62199F50FFC1FFE5FF34FB40FD03F834	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
62199F50FFCDFFE0FF34FA91FE6AFC99.text	62199F50FFCDFFE0FF34FA91FE6AFC99.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Limnonectes gyrinophorus Dehling & Neokleous & Das & Grafe & Min & Hertwig 2025	<div><p>Limnonectes gyrinophorus sp. nov.</p><p>Rana finchi — Inger &amp; Stuebing 1992: 46 (partim); Inger et al. 1996: 363; Malkmus 1996: 293; Lakim et al. 1999: 32. Limnonectes finchi — Inger et al. 2000: 16 (partim); Hoffmann 2002: 28, 31; Malkmus et al. 2002: 141 (partim); Inger et al.</p><p>2017: 79 (partim).</p><p>Holotype. NMBE 1075255, adult female, from Sungai Kipungit 1 (6.051250°, 116.701919°; 530 m), Kinabalu National Park, Poring, Sabah, Malaysia collected on 11 March 2017 by A. Haas, R. Hagmann, S. T. Hertwig, P. Yambun (Fig. 3J, Fig. 12).</p><p>Paratype. ZFMK 79844, adult male, from Poring, Sabah, Malaysia, collected on 16 April 1994 by Peter Hoffmann .</p><p>Diagnosis. Limnonectes gyrinophorus sp. nov. belongs to the L. finchi species group, showing the typical characters of the group, including moderate size (SVL of males to 36.4 mm, of females to 42.9 mm); odontoid processes in lower jaw low and inconspicuous in both males and females; dorsum granular with interrupted, longitudinal dorsal rows of elongate tubercles; interorbital distance shorter than upper eyelid width; pineal spot distinctly visible; toes not fully webbed; and horseshoe-shaped dermal ridge in middle of dorsum posterior to arm insertion. The species differs from the other species of the group by showing a unique combination of characters including dorsal face very rough with many elevated warts and glandular ridges, and cloaca surrounded by numerous, densely arranged small tubercles; skin on ventral side of head and chest thick, forming collar-like posterior edge and distinct transverse chest fold; loreal region angular; pineal spot round, in front of line between anterior eye edge; vomer processes bearing 3–4 teeth, separated from each other and from choana by half length of individual process; hindlimbs relatively short with tibiotarsal joint reaching to nostril; first finger relatively long, subequal to third finger; distinct, moveable dermal flap on postaxial side of Toe V from proximal end of metacarpus to disk. The species differs from the other species of the group by at least 4.4% in the sequence of the 16S barcoding gene.</p><p>Description of holotype. Adult female; SVL 42.9 mm; body sturdy, widest at level of arm insertion, markedly tapering to groin (Fig. 3J); head large (HL/SVL 0.38, HW/SVL 0.38), about as wide as long (HW/HL 0.98); snout moderately long (SL/HL 0.42), rounded in both dorsal view and profile, slightly projecting beyond lower jaw, slightly wider than long (SL/EE 0.95); canthus rostralis distinct between eye and nostril, concave in anterior half, straight-lined posteriorly; loreal region oblique, moderately concave; nostrils rounded, directed dorsolaterally, situated closer to tip of snout than to eye (EN/NS 1.23), separated from each other by distance slightly larger than distance between eye and nostril (NN/EN 1.10); eye directed anterolaterally, moderately protruding, large (ED/HL 0.31), its diameter shorter than snout (ED/SL 0.73); interorbital distance shorter than upper eyelid width (IO/EW 0.81) and smaller than internarial distance (IO/NN 0.73); pineal spot distinctly visible, rounded, anterior to line between anterior eye edges; tympanum and its annulus distinctly visible; tympanum separated from eye by threefifths its diameter (ET/TD 0.59); tympanum diameter smaller than eye diameter (TD/ED 0.63); upper jaw with dentition; odontoid processes in lower jaw low, inconspicuous; choanae small, rounded, located anterolaterally at margins of roof of mouth, not covered by palatal shelf of maxilla in ventral view; vomer processes bearing teeth, large, separated from each other and from choana by half length of individual process (Fig. 4J); tongue long and broad, bilobed for about one-fifth of its length, free distally for about two-thirds its length; median lingual process absent.</p><p>Dorsal surfaces of head, trunk and limbs rough, with scattered enlarged tubercles, larger ones with pointed keratinous tips, and with elongate short dermal ridges (Fig. 12); tubercle ridges in indistinct, interrupted longitudinal rows on both sides parallel to middle of the dorsum, replaced by indistinct, inverted-v-shaped dermal ridge in middle of back just behind level of arms; dorsal surfaces of forearm, hand, tarsus and foot shagreened; lateral surfaces of trunk shagreened with scattered small tubercles; thick dorsal skin of trunk ending abruptly at level of posterior end of upper eyelid, forming distinct transverse dermal edge; dorsal face of head finely granular; skin from nostril to tympanum rougher granular; supratympanic fold very thick and conspicuous, extending from posterior end of orbit to insertion of arm, curved; enlarged tubercles on posterior part of upper eyelid; very conspicuous and large tubercle just posterior to mouth opening; skin on ventral side of head smooth with distinct wrinkles; skin smooth on chest and abdomen; skin on ventral side of head and chest thick, forming collar-like posterior edge running from end of jaw, anteriorly to arm insertion to middle of chest, eventually forming distinct transverse chest fold between arm insertions; ventral side of limbs smooth; cloaca without dermal flap, surrounded by numerous, densely arranged small tubercles.</p><p>Forelimbs moderately sturdy; hand relatively small (HND/SVL 0.24); tips of fingers rounded, swollen and enlarged into small disks, largest on Finger I, smallest on Finger IV; relative length of fingers: IV &lt;II &lt;I &lt;III, with Finger I subequal to Finger III, and Finger IV subequal to Finger II; subarticular tubercles rounded, well developed, numbering one on Fingers I and II, two on Fingers III and IV, proximal tubercles on Fingers III and IV much larger and more prominent than distal ones; subarticular tubercle of Finger I markedly larger than subarticular tubercle on Finger II and proximal tubercles on Fingers III and IV; finger webbing absent; thenar tubercle distinct, oval, flat, about one-third length of metacarpal of Finger I; inner palmar tubercle small, rounded, flat, on proximal end of metacarpus of Fingers II and III; outer palmar tubercle larger, elongated, on proximal half of metacarpal of Finger IV, fused with inner palmar at proximal base, about two-thirds size of thenar tubercle.</p><p>Hindlimbs moderately sturdy, long (LEG /SVL 1.72); heel reaching to level of nostrils when legs adpressed forwardly to body; tibiofibula long (TFL/SVL 0.55), slightly longer than thigh (TFL/THL 1.10); heels overlapping each other considerably when knees flexed and thighs held perpendicularly to median plane; foot shorter than tibiofibula (FOT/TFL 0.89); relative length of toes: I &lt;II &lt;V &lt;III &lt;IV; toe tips rounded, slightly enlarged into disks; subarticular tubercles numbering one on Toes I and II, two on Toes III and V, and three on Toe IV; pedal webbing formula I 1+/2 II 1+/2.25 III 1.25/2.75 IV 2.75/1.25 V (Fig. 6); dermal flap on postaxial side of Toe V, widest from proximal end of metacarpus to proximal subarticular tubercle, continued to disk; inner metatarsal tubercle very prominent, elongated, half as long as metatarsus of Toe I; outer metatarsal tubercle absent; low ridges of callous tissue from proximal plantar part of metatarsus to proximal subarticular tubercle on Toes I, II, and V.</p><p>Colouration. Life colouration (Fig. 12): Dorsum medium greyish brown; larger tubercles light to reddish brown; interorbital bar dark brown, continued onto posterior part of upper eyelids, bordered anteriorly by light brown band; dorsal face of head anterior to interorbital bar light olive-greenish brown with dark brown speckling; pineal spot whitish; sides of head to tympanic area with alternate bands of dark olive-greenish brown and greenish white bands; tympanum covered with dark brown spot; supratympanic fold light brown; flanks light greenish grey; arms creamy grey with faint greyish crossbars on lower arm; fingers light cream-coloured with dark grey blotches; legs greenish light brown with olive-greenish brown crossbars and small, light reddish brown tubercles; crossbars as wide on thigh as on crus; toes light cream-coloured; webbing dark olive green; ventral side of head, arms, and legs off-white; dark brown spots along mandibular edge; throat mottled dark brown. See Hoffmann (2002: 28) and Malkmus et al. (2002: 141) for a photo of the male paratype in life.</p><p>In preserved state, dorsum dark greyish brown; dorsal pattern elements dark brown (Fig. 3J); limbs including fingers and toes somewhat lighter brown, their reddish life colouration faded to light brown; ventral side light grey.</p><p>Variation. The male paratype matches the holotype in general appearance. SVL of the adult male is 36.4 mm (Table 2). The male lacks nuptial pads and has numerous subconcentric dermal folds on the ventral side of the head. The vocal sac aperture is slit-like, directed posterolaterally, situated at corner of mouth. Pedal webbing variation is I 1+/2 II 1 + [50],1.25 [50] /2.25 [50],2.5 [50] III 1.25 /2.75 [50],3- [50] IV 2.75 [50],3+ [50] /1.25 V (Fig. 6) .</p><p>Bioacoustics. The call was reported by Malkmus et al. (2002) to consist of a short trill, but it is unclear if this characterization was based on a direct observation of a specimen from Kinabalu (Poring) or if the description of the call of L. finchi by Inger &amp; Stuebing (1989) was cited. No call recording was available for a detailed analysis.</p><p>Distribution. So far, the species is known only from the eastern flank of Gunung Kinabalu in the area of Poring Hot Springs in northern Sabah, Malaysia at elevations of 500–530 m (Figs. 8, 9).</p><p>Ecology. The male paratype (ZFMK 79844) was observed near Poring carrying seven tadpoles to an ephemeral pond next to a road along the boundary of the Kinabalu National Park (Hoffmann 1998, Malkmus et al. 2002). An uncollected male carrying at least 30 tadpoles on its back was observed north of Poring on 1 December 2024 (H. P. Guek in in litt., Fig. 10H).</p><p>Etymology. The species epithet is composed of the Ancient Greek noun γυρίνος [gyrínos], meaning tadpole, and the Latinized suffix -phorus (“the one who carries”; from Ancient Greek φορεῖν [phoreĩn] = “to carry”); meaning “the one that carries the tadpoles”, in allusion to the tadpole-carrying behaviour of the males of the species.</p><p>Suggested English name. Rugged Guardian Frog.</p><p>Species delimitations</p></div>	https://treatment.plazi.org/id/62199F50FFCDFFE0FF34FA91FE6AFC99	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.		MagnoliaPress via Plazi	Dehling, Maximilian;Neokleous, Dario N.;Das, Indraneil;Grafe, Ulmar;Min, Pui Yong;Hertwig, Stefan T.	Dehling, Maximilian, Neokleous, Dario N., Das, Indraneil, Grafe, Ulmar, Min, Pui Yong, Hertwig, Stefan T. (2025): Cryptic radiation within the tadpole-carrying Guardian Frogs from Borneo, Limnonectes palavanensis and L. finchi (Anura: Dicroglossidae), with the description of eight new species. Zootaxa 5650 (1): 1-80, DOI: 10.11646/zootaxa.5650.1.1, URL: https://doi.org/10.11646/zootaxa.5650.1.1
