Kalophrynus misbahulmuniri, Gonggoli & Fauzan & Kaprawi & Herlambang & Kirono & Wiradarma & Alif & Hamidy, 2025

Gonggoli, Ade Damara, Fauzan, Muhammad Fakhri, Kaprawi, Fajar, Herlambang, Alamsyah Elang Nusa, Kirono, Sasi, Wiradarma, Huda, Alif, Haegal & Hamidy, Amir, 2025, Taxonomic assessment of Javanese Kalophrynus Tschudi, 1838, with a description of a new species (Amphibia, Anura, Microhylidae), Zootaxa 5646 (4), pp. 501-526 : 513-521

Kalophrynus misbahulmuniri sp. nov.

( Figs. 8–10 , 11A , 13 )

K. pleurostigma : Iskandar (1998)

K. minusculus : Kurniawan et al. (2022), Kaprawi et al. (2022a, b)

Holotype. Adult male, MZB Amph 32726, collected from Sagara Village , Cibalong District, Garut Regency, West Java Province, Indonesia (-7.700111, 107.838722, 14 m a.s.l), by Misbahul Munir, Ade Damara Gonggoli, Fajar Kaprawi, and Haegal Alif on 4 April 2022.

Paratypes (n=11). Three males ( MZB Amph 32720, 32724–25) and three females ( MZB Amph 32721–23) were collected together with the holotype . One male ( MZB Amph 32695) and two females ( MZB Amph 32693–94) from Sagara Village , Cibalong District, Garut Regency, West Java Province, Indonesia, collected by Umar Fadli Kennedi, Mohammad Ali Ridha, Dzikri Ibnul Qayyim, and Rizky Rafsazan in April 2017 . One male ( MZB Amph 504) from Nusa Kambangan Island, Cilacap Regency, Central Java Province, Indonesia, collected by S.S. Liem on 8 November 1962 . One male ( MZB Amph 31927) from Tambakreja, Nusa Kambangan Island, South Cilacap District, Cilacap Regency, Central Java Province, Indonesia (-7.728713, 108.8149, 96 m a.s.l) collected by Wahyu Sigit, Frendi Irawan, Diagal Wisnu, Hening Triandika Rachman, Nanang Kamaludin, and Nukman, on 15 September 2020 .

Etymology. The specific epithet “misbahulmuniri ” is an eponym dedicated to the late Misbahul Munir, who contributed significantly to the taxonomy and conservation of Indonesian herpetofauna and was one of the type specimen collectors.

Suggested English common name. Misbahul Munir’s Sticky Frog.

Suggested Indonesian common name. Katak lengket Misbahul Munir.

Diagnosis. The new species is assigned to Kalophrynus based on molecular analyses and the following combination of morphological characters: absence of spine-like projection of skin at heel and elbow; fingers without greatly enlarged tubercles; snout less than twice the diameter of the eye; inner metatarsal tubercle low, not shovel-like; tympanum visible ( Inger 1966).

The new species can be distinguished from its congeners by the following combination of characters: medium size (SVL 26.9–31.8 mm in adult males, 32.8–38.5 mm in adult females); inguinal spot usually present, with light border; hand and foot with distinct subarticular tubercles; hindlimb moderate length; outer palmar tubercle large; webbing on fourth toe reaches middle subarticular tubercle or slightly less; thin dorsolateral line; absence faint hourglass mark and conical spines on the dorsal skin of the body.

Description of holotype. Adult male ( Fig. 8A–E , 10A ); medium-sized (SVL 31.8 mm); habitus stocky; head broader (HW 10.2 mm, 32.0 % SVL) than long (HL 8.9 mm, 28.1 % SVL); snout sharply pointed in lateral view, protruding beyond lower jaw; eye diameter (ED 3.9 mm, 12.1 % SVL) longer than snout length (SL 3.5 mm, 10.9 % SVL); pupil horizontal; canthus rostralis distinct; loreal region vertical, slightly convex; nostril below canthus, nearer to tip of snout (SNL 1.0 mm, 3.0 % SVL) than to eye (NEL 2.4 mm, 7.6 % SVL); interorbital distance (IOD 4.2 mm, 13.2 % SVL) wider than internarial distance (IND 2.0 mm, 6.1 % SVL), the latter nearly equal to upper eyelid (UEW 2.1 mm, 6.6 % SVL); pineal spot absent; tympanum distinct, roughly rounded, diameter (TD 2.6 mm, 8.3 % SVL) about two-thirds of eye; upper jaw edentate; tongue smooth, without papillae; vomerine fold smooth, straight-edged flaps behind choanae, each extending from labial edge of mouth to near midline, separated by short gap from its opposite fold; postorbital low, smooth; buccal fold distinct, higher than postorbital; vocal slit present, on both side of mouth floor between base of tongue and angle of jaws; median subgular vocal sac.

Forelimb slender; finger webbing absent; fingertips blunt, rounded; finger length formula IV <I <II <III, fourth finger (Fin4L 1.8 mm, 5.8 % SVL) slightly shorter than first (Fin1L 2.0 mm, 6.3 % SVL); portion of fourth finger projection from palm longer than terminal phalanx of third finger; subarticular tubercles on fingers and palmar tubercles well-developed, rounded, two on fingers I, II, and IV, three on III; outer palmar tubercle large, oval; humeral spine absent; nuptial pad absent.

Hindlimb slender, moderately long (HLL 47.6 mm, 149.8 % SVL); toe length formula I <II <V <III <IV; fifth toe projecting as far as third toe; foot (FL 12.2 mm, 38.2 % SVL) shorter than tibia (TBL 12.9 mm, 40.6 % SVL); tibio-tarsal articulation of adpressed limb reaching behind tympanum; toe tips blunt, rounded; toes webbing poorly developed, webbing formula I 1–1 II 1–2 III 1–3 IV 3.5–1 V; fifth toe projecting slightly below the third toe; subarticular tubercles rounded, one on toes I and II, two on III and V, three on IV; inner metatarsal distinct, oval; outer metatarsal present, indistinct.

Dorsum skin finely granular, with few small whitish tubercles, scattered from snout to vent; dorsolateral lined with small whitish tubercles, extended from tip of snout to groin; gland on side of head, behind tympanum, delimited by sinuous groove; supratympanic fold curved; abdomen, ventral thigh, and cloaca with large flattened glandules; pair of round, whitish, slightly elevated tubercles on pectoral region.

Coloration. In life, dorsum yellowish brown, with a very indistinct pair of dark bands from between the eyes to the groin; whitish tubercles formed a thin dorsolateral line from the anterior upper eyelid to the groin, forming the boundary between lighter dorsum and darker side of head and flank; dorsal limbs with dark cross bars; inguinal region with rounded black spot, with whitish tubercle edges; chin and pectoral region brownish; abdomen and ventral thigh cream. In preservatives, body coloration fades to dark brown, and the side of the body to blackness; chin and pectoral region fade to dark gray; ventral of the thigh and abdomen fades to cream.

Variation. Morphometric variations among the specimens show that males have larger in RHW, RICD, RIOD, Rfin2L, Rfin3L, RFML, and RToe5L than females, while females have larger in SVL, RSNL, RUEW, RToe1L, and RToe2L than males ( Table 2). In life, dorsal body coloration varies from light peach ( Fig.9B, D, E, F ) to burnt orange ( Fig. 9C ). A female paratype ( MZB Amph 32722) exhibited light peach coloration in life ( Fig. 9E ), which changed to dark brown after euthanasia ( Fig. 10B 1 ). The dorsal skin of a male paratype ( MZB Amph 31927) was light grayish with scattered black spots and several dark stripes near the groin ( Fig. 10E 1 , E 3 ). A male paratype ( MZB Amph 32720) has a deformed right foot ( Fig. 10A 2 ). Among the specimens, an indistinct pair of dark bands on the dorsal skin of the body was observed in one female ( MZB Amph 32723). The inguinal spot was absent in one female ( MZB Amph 32693). In most paratype specimens, the proximal subarticular tubercle on the fifth toe was indistinct. Three female paratypes had a variation in toe webbing: one female ( MZB Amph 32693) was a little bit dried and showed poorly developed webbing on the fourth toe, webbing formula: I 1–2 II 1.5–3 III 1.5–3.5 IV 3.5–1.5 V, while two females ( MZB Amph 32721, 32723) had webbing formula: I 1–2 II 1–3 III 1.5–3.25 IV 3.5–1.5 V. Fourth toe of all females projecting as far as third .

Morphological comparison. Kalophrynus misbahulmuniri sp. nov. is easily distinguished from K. minusculus by more developed toe webbing, I 1–1 II 1–2 III 1–3 IV 3–1 V (vs. I 1–2 II 1–3 III 2–4 IV 4–2 V); a pair of black bands and stripes on dorsal skin of body absent and indistinct (vs. present and distinct); snout sharply pointed (vs. rounded); larger body size in mean values of adult males, SVL 26.9–31.8 mm, mean 29.6 ± SD 1.6 (vs. 23.1–29.2 mm, mean 26.0 ± SD 1.7), and adult females, SVL 32.8–38.5 mm, mean 36.1 ± SD 2.4 (vs. 29.3–32.3 mm, mean 30.8 ± SD 1.5).

Kalophrynus misbahulmuniri sp. nov. differs from K. pleurostigma by females having smaller body size in mean values, SVL 32.8–38.5 mm, mean 36.1 mm ± SD 2.4 (vs. 34.5–43.5 mm, mean 40.9 mm ± SD 3.2), thinner dorsolateral line, Fig. 11A (vs. thicker, Fig. 11C, D ), fifth toe projecting as far as the third toe in females, Fig. 12A (vs. fifth toe not projecting as far as third, Fig. 12C ), and absence of hourglass marking on dorsal skin (vs. present in some specimens). The new species differs from K. meizon by having a smaller body size, SVL 26.9–31.8 mm in males, 32.8–38.5 mm in females (vs. 37.0–50.0 mm in males, 35.0–60.0 mm in females), and absence of conical spines on the dorsal skin of the body of males [vs. present: Zug (2015)].

Kalophrynus misbahulmuniri sp. nov. is easily differentiated from the nubicola group ( K. nubicola , K. puncak , K. dringi ) by having larger body size, SVL 26.9–31.8 mm in males, 32.8–38.5 mm in females (vs. ≤ 22.7 mm in males, ≤ 24.8 mm in females), distinct subarticular tubercles on hand and foot [vs. indistinct: Fukuyama et al. (2021)], distinct snout projection [vs. indistinct: Fukuyama et al. (2021)], and dorsolateral line present [vs. absent (except K. dringi ): Fukuyama et al. (2021)].

Kalophrynus misbahulmuniri sp. nov. differs from K. barioensis , K. bunguranus , K. cryptophonus , K. eok , K. robinsoni , K. subterrestris , and K. tiomanensis by having a larger body size, SVL 26.9–31.8 mm in males, 32.8–38.5 mm in females [vs. ≤ 26.3 mm in males, ≤ 27.0 mm in females (except in females K. cryptophonus )]. The new species differs from K. anya , K. baluensis , K. interlineatus , K. kiewi , K. orangensis , and K. sinensis by having a smaller body size, SVL 26.9–31.8 mm in males, 32.8–38.5 mm in females [vs. ≥ 33.7 mm in males, ≥ 39.4 mm in females (except in females K. anya , K. interlineatus , K. kiewi , and K. orangensis )].

Kalophrynus misbahulmuniri sp. nov. has an overlapping body size with K. calciphilus , K. intermedius , K. palmatissimus , and K. punctatus , but can be differentiated from those species by the presence of inguinal spot in most specimens (vs. absent).Additionally, the new species differs from K. calciphilus by having smaller subarticular tubercles on the hand and foot [vs. larger: Dehling (2011)]. From K. intermedius , the new species differs by the fifth toe projecting as far as the third toe in females (vs. fifth toe not projecting as far as third). From K. palmatissimus and K. punctatus , the new species differs from those species by webbing on the fourth toe usually reaching the middle subarticular tubercle or slightly less (vs. webbing on the fourth toe webbed to the distal subarticular tubercle in K. palmatissimus and below the proximal subarticular tubercle in K. punctatus ).

The new species also has an overlapping body size with K. heterochirus , K. honbaensis , K. limbooliati , and K. yongi , but, respectively, the new species can be differentiated from these species as follows: from K. heterochirus by the presence of black inguinal spot (vs. white-bluish) and portion of fourth finger projection from palm longer than terminal phalange of third finger (vs. shorter); from K. limbooliati by the portion of fourth finger projection from palm longer than terminal phalange of third finger (vs. shorter); from K. yongi by the absence of nuptial pad (vs. present); and from K. honbaensis by the absence of finger webbing (vs. present).

Call characteristics. A male paratype ( MZB Amph 32720) was collected from the field and brought to a room. Six calls of the specimen were recorded at distances of approximately 0.5–1 m at an air temperature of 24.0°C on 22 April 2022. Call consists of two types of pulsed note series ( Fig. 13A ), sounding to the human ear like “krek…. krek….krek... krek..krek..krek..krek..krek” with duration of call 1.56− 13.89 s (mean 5.066 ± SD 4.645), inter call interval 0.920 –1.681 s (mean 1.262 ± SD 0.386), and dominant frequency 1378–2756 Hz (mean 1600 ± SD 322.402). The interval between the two types of note series was 0.753 –12.663 s (mean 3.600 ± SD 5.120) .

Note series I ( Fig. 13B ) consist of 1–3 notes, with 2–6 pulses, duration of note series 0.023 −1.998 s (mean 0.471 ± SD 0.773), pulse duration 0.005 −0.038 s (mean 0.017 ± SD 0.008), inter-pulse interval 0.000 −0.018 s (mean 0.002 ± SD 0.005), pulse period 0.010 −0.026 s (mean 0.016 ± SD 0.004), pulse rate 39.141 −103.041 s (mean 67.886 ± SD 16.452), and dominant frequency 1378.125−2239.453 Hz (mean 1531.25 ± SD 138.29).

Note series II ( Fig. 13C ) consists of 3−9 notes, with 2–7 pulses, duration of note series 0.377 −2.541 s (mean 1.37 ± SD 0.937), pulse duration 0.004 −0.037 s (mean 0.015 ± SD 0.007), inter-pulse interval 0.000 −0.017 s (mean 0.002 ± SD 0.004), pulse period 0.008 −0.026 s (mean 0.015 ±SD 0.003), pulse rate 38.458 −119.357 s (mean 68.538 ± SD 15.133), and dominant frequency 1378.125−2756.250 Hz (mean 1618.764 ±SD 364.146).

Call comparisons. The call of Kalophrynus misbahulmuniri sp. nov. differs from K. baluensis , K. barioensis , K. limbooliati , K. meizon , and K. yongi by the well-pulsed note (vs. unpulsed note). The call of new species differs from K. minusculus by consisting of 2−7 pulses (vs. only two pulses), and a higher dominant frequency, 1378−2239 Hz (vs. 516–689 Hz). The call of new species differs from K. cryptophonus , K. dringi , K. interlineatus , K. nubicola , and K. puncak by consisting of a series of notes (vs. a single note). Additionally, the call of new species differs from K. cryptophonus and K. interlineatus by having a higher dominant frequency, 1378−2239 Hz (vs. 900−1200 in K. cryptophonus , 660−820 Hz in K. interlineatus ). The call of new species differs from K. calciphilus by having two types of note series (vs. a single type of note series) and consists of 2−7 pulses (vs. only consisting of 7 or 8 pulses).

Tadpoles. Unknown.

Distribution and natural history. Kalophrynus misbahulmuniri sp. nov. is known only from Sagara and Nusa Kambangan Island at a low elevation between 14−19 m a.s.l. Holotype and paratypes from Sagara were found on leaf litter in secondary forests at low elevations near coastal areas ( Fig. 14 ). This new species was also recorded in Leuweung Sancang ( Kaprawi et al. 2022a, b). The tadpole is unknown. The anuran species had been found sympatrically with the new species: Chirixalus pantaiselatan Munir, Hamidy, Kusrini, Kennedi, Ridha, Qayyim, Rafsanzani & Nishikawa , Fejervarya limnocharis (Gravenhorst) , Hylarana nicobariensis Stoliczka , Kaloula baleata (Müller) , Limnonectes macrodon (Duméril & Bibron) , Limnonectes microdiscus (Boettger) , Microhyla achatina Tschudi , Microhyla palmipes Boulenger , and Polypedates leucomystax (Gravenhorst) .

Figures

FIGURE 1. Map showing known localities of the Javanese Kalophrynus pleurostigma (triangle), K. minusculus (star), K. pleurostigma “Sumatra II” (round). 1) Nusa Kambangan; 2) Sagara and Leuweung Sancang; 3) Ujung Kulon; 4) Gelugur; 5) Sedayu. The land topography was generated using the SRTM 90 m dataset from CGIAR-CSI (https://srtm.csi.cgiar.org), processed with QGIS 3.22 Białowieża, and visualized in Blender 3.6.2 (https://www.blender.org).

FIGURE 2. Bayesian tree from a 487 bp sequence of mitochondrial 16S rRNA genes for Kalophrynus species and related species. Numbers near branches represent bootstrap support for Bayesian Inference posterior probabilities (PP) and Ultrafast Bootstrap Probability (UFB).

FIGURE 3. Scatterplot of the first (PC1) and second (PC2) principal components shows the clustering of morphological measurements of males (A) and females (B) among Javanese Kalophrynus pleurostigma (yellow), K. minusculus (red), K. pleurostigma (green), and K. meizon (blue).

FIGURE 8. Dorsolateral view, dorsal view, ventral view, lateral view, ventral view of the right hand, and ventral view of the right foot, respectively, of the adult male holotype of Kalophrynus misbahulmuniri sp. nov., MZB Amph 32726 (A, C–H), and the adult female paratype, MZB Amph 32721 (B, J–O); ventral view of head with opened mouth of holotype (I), showing the buccal and postorbital fold. Scale bar = 5 mm. Photographs by Misbahul Munir.

FIGURE 9. Dorsolateral view of Kalophrynus misbahulmuniri sp. nov. in life. Adult male paratype MZB Amph 32720, and adult female paratype, MZB Amph 32721 in amplexus position (A); adult male paratype (MZB Amph 32724); adult male, MZB Amph 32720 (C); adult female paratype, MZB Amph 32721 (D); adult female, MZB Amph 32722 (E); adult female, MZB Amph 32723 (F). Photographs by Misbahul Munir.

FIGURE10.Dorsalview, ventralview, lateralview,ventralview of hand, andventralview of footofKalophrynus misbahulmuniri sp. nov. Adult male paratype, MZB Amph 32720 (A1–A5); adult female paratype, MZB Amph 32722 (B1–B5); adult female paratype, MZB Amph 32723 (C1–C5); adult male paratype, MZB Amph 31924 (D1–D5); adult male paratype, MZB Amph 31927 (E1–E5); adult female paratype, MZB Amph 32693 (F1–F4); adult male paratype, MZB Amph 504 (G1–G4); dorsal view and ventral view of adult female, MZB Amph 32694 (H1–H2), and adult male, MZB Amph 32695 (I1–I2). Scale bar = 5 mm. Photographs by Misbahul Munir.

FIGURE 11. Adult male of Kalophrynus misbahulmuniri sp. nov., MZB Amph 32726 (A); adult male of K. minusculus, MZB Amph 27229 (B); adult male of K. pleurostigma from PuncakAnai, MZB Amph 32288 (C); and adult female of K. pleurostigma from Muara Bangko, MZB Amph 32252 (D), in life. Photographs by Misbahul Munir (A, C), Sasi Kirono (B), and Fajar Kaprawi (D).

FIGURE 12. Ventral view of right foot of adult female Kalophrynus misbahulmuniri sp. nov., MZB Amph 32723 (A), adult female K. minusculus, MZB Amph 27234 (B), and adult female K. pleurostigma, KUHE 63828 (C), showing the fifth toe projecting against the third toe. Scale bar = 5 mm. Photographs by Misbahul Munir.

FIGURE 13. Oscillogram and spectrogram of one call (A) of adult male paratype of Kalophrynus misbahulmuniri sp. nov., MZB Amph 32720, which consists of note series I with three notes (B), and note series II with nine notes (C).

FIGURE 14. Natural habitat of Kalophrynus misbahulmuniri sp. nov. in Sagara village. Photograph by Misbahul Munir.