Megophrys lancip, Munir & Hamidy & Farajallah & Smith, 2018

Munir, Misbahul, Hamidy, Amir, Farajallah, Achmad & Smith, Eric N., 2018, A New Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae) from southwestern Sumatra, Indonesia, Zootaxa 4442 (3), pp. 389-412 : 395-408

Megophrys lancip sp. nov.

( Figs. 3–5 )

Holotype. MZB Amph 22233 (field number ENS 14509) an adult male from the north-east of Desa Ngarip, Kecamatan Ulubelu, Kabupaten Tanggamus, Province of Lampung, Sumatra (5.31359°S, 104.53151°E, elevation 1010 m. a.s.l.), collected by Eric Nelson Smith, Amir Hamidy and Elijah Wostl, at 2005 h on 10 June 2013 ( Fig. 3 , 4A–E ).

Paratypes. UTA A-64763 (field number ENS 7579 ; formerly MZB Amph 12564) adult male from the Bengkulu to Kepahiang road, Kecamatan Taba penanjung, Kabupaten Bengkulu Tengah, Province of Bengkulu, Sumatra (3.68561°S, 102.53861°E, 620 m. a.s.l.), collected by Eric N. Smith and Michael B. Harvey, at 1740 h on 30 May 1996 ( Fig. 5B ) ; MZB Amph 3469, 14602, and 26738 ( Fig. 4F–J ), three adult females from Bukit Barisan Selatan National Park, at Desa Kubu Perahu , near Kota Liwa , Kecamatan Balik Bukit , Kabupaten Lampung Barat, Province of Lampung, Sumatra, collected by Andiek Fajar on 19 February 1998 (5.07870°S, 104.01189°E, 434 m. a.s.l.), by Inggar Ul-Hasanah on 17 September 2005 (5.07526°S, 104.04081°E, 550 m. a.s.l.), and by Anton Nurcahyo in 2005 (5.07526°S, 104.04081°E, 550 m. a.s.l.), respectively ; MZB Amph 3 472 adult female from Bukit Barisan Selatan National Park, at Desa Sukaraja , Kecamatan Semaka , Kabupaten Tanggamus , Province of Lampung, Sumatra (5.51955°S, 104.45303°E, 526 m. a.s.l.), collected by Andiek Fajar on 6 November 1997 .

Etymology. The species name lancip is the Indonesian word for “pointed”, used as an adjective, and reflects the extremely pointed rostral appendage of the new species.

Suggested English common name. Pointed Horned Frog.

Suggested Indonesian common name. Katak-tanduk lancip .

Diagnosis. The new species is assigned to the genus and subgenus Megophrys based on our molecular analysis ( Fig. 2 ) and the following morphological features: (1) absence of a distinct glandular fold forming a Y, X, or H on the parietoscapular region, possession of (2) a broad and depressed head, (3) pectoral and femoral glands, (4) eyelid appendages, (5) a vertical pupil, (6) a pair of conical spine at the corner of the mouth, and (7) a protruding snout ( Delorme et al. 2006). Megophrys lancip sp. nov. is a medium-sized Megophrys (adult males SVLh 37.9–47.7 mm, n = 2; adult females SVLh 38.7–82.5 mm, n = 4) with the head shorter than the body (adult males HL 15.3–19.9 mm: 40.3–41.7% of SVLh, n = 2; adult females HL 16.0– 35.4 mm: 41.4–42.9% of SVLh, n = 4), and broader compared to its length (adult males HL 87.7–95.8% of HW, n = 2; adult females 80.8–85.7% of HW, n = 4). The snout is moderate length (adult males SL 30.5–35.8% of HL, n = 2; adult females SL 29.4–33.6% of HL, n = 4), extremely pointed in dorsal view, protruding in lateral view, and with a sharp and pointed (acuminate) rostral appendage that projects beyond the lower jaw. The rostral appendage is moderate (adult males SNLh 3.0– 3.3 mm, 7.0–8.0% to SVLh, n = 2; adult females SNLh 2.5–5.4 mm, 4.9–6.7% of SVLh, n = 4; adult males SAL; 1.2–1.4 mm, 2.9–3.1% of SVLh, n = 2; adult females SAL 0.6–3.44 mm, 1.5–4.1% of SVLh, n = 4); the upper eyelid has a triangular, short, and narrow-based appendage; the tympanum is distinct, small, oval, andoriented vertically; vomerine teeth are present; and the subarticular tubercle and outer metatarsal tubercles are absent.

Description of Holotype (measurements in mm). Adult male; body stocky, robust (SVL 36.8; SVLh 37.9); head broad and depressed (HW 17.4: 46.0% of SVLh), slightly broader than long (HL 15.3: 40.3% of SVLh); snout extremely pointed from dorsal view, protruding in lateral view, tip with an acuminate sharp rostral appendage projection (SAL 1.2: 3.1% of SVLh), projecting beyond lower jaw; eye large and lateral on head, smaller (ED 5.0: 13.1% of SVLh) than snout length (SL 5.5: 14.4% of SVLh), eye diameter about 3.5 times of tympanum diameter (ED 5.0: 359.4% of TDH); canthus rostralis sharp, angular, concave; nostril below canthus on lateral view, distinctly closer to eye (NEL 1.9: 4.9% of SVLh) than to snout (SNL 2.3: 6.0% of SVLh); internarial distance (IND 3.4: 8.9% of SVLh) much shorter than interorbital distance (IOD 5.1: 13.5% of SVLh); upper eyelid with low tubercles and short narrow-base eyelid appendage projection (EHL 1.2: 2.9% of SVLh) much shorter than total width (UEWh 4.8: 12.7% of SVLh); a conical spine present at corner of mouth; tympanum distinct, oval, oriented vertically (TDV 2.2: 5.7% of SVLh; TDH 1.4: 3.6% of SVLh); vomerine teeth present; tongue not notched, without papilla.

Forelimbs slender and short (FLL 25.5: 67.3% of SVLh), about half of hindlimbs (FLL 25.5: 49.0% of HLL); fingers moderately slender, unwebbed, without lateral fringes, with rounded and swollen tips; first finger (fin1L 3.1: 8.1% of SVLh) slightly longer than second (fin2L 3.0: 7.8% of SVLh) and fourth (fin4L 2.8: 7.5% of SVLh), third much longer (fin3L 5.0: 13.1% of SVLh); finger length formula IV<II<I<III; subarticular tubercle absent; inner palmar tubercle (IPTL 3.6: 9.4% of SVLh) larger than outer palmar tubercle (OPTL 1.2: 3.2% of SVLh), inner palmar tubercle extending from base to first subarticular tubercle of first finger ( Fig. 4 ). Hind limb slender and relatively long (HLL 52.0: 137.2% of SVLh); tibia (TL 16.0: 42.1% of SVLh) longer than tarsus (TSL 9.1: 23.9% of SVLh) and slightly longer than foot (FL 15.0: 39.7% of SVLh); tips of toes similar to those of fingers, rounded and swollen; first toe (Toe1L 1.9: 4.9% of SVLh) much shorter than second (Toe2L 2.4: 6.4% of SVLh), third (Toe3L 4.9: 12.9% of SVLh), and fifth (Toe5L 2.6: 6.9% of SVLh), and fourth toe longest (Toe4L 7.3: 19.3% of SVLh), toe length formula I<II<V<III<IV; tibio-tarsal articulation, of depressed hind limb towards body, reaches middle of eye; toe subarticular and outer metatarsal tubercles absent, inner metatarsal tubercle smaller than inner palmar tubercle (IMTL 2.5: 69.5% of IPTL); toes webbed only at base, except for fifth toe, webbing formula, I (1), II (1–1), III (2–2), IV (3–3), V (1 1/2).

Skin. Dorsal skin is smooth with low, sharply angular, tubercles from the snout to the vent, especially around the waist; the tubercles are denser laterally on the body and on the posterior surface of the thigh; upper eyelids with low conical tubercles; supratympanic fold distinct, angular, widening anteriorly, narrow posteriorly, extending from posterior of eye, curving down around upper border of tympanum and terminating above axilla; Y, X or H fold absent between parietoscapular area to level of axilla; dorsolateral fold extends from between eye and tympanum to the groin, wider from shoulder to groin; forelimb and hindlimb surface tuberculate and with transverse folds in forearm, tibia, and femur; a weakly developed tubercles ridge on outer of edge of forearm and tarsus forming a weak serration; ventral skin slightly tuberculate; skin on throat wrinkled, chest slightly tuberculate; ventral surface of limbs and abdomen smooth to weakly granular; a pair of white pectoral glands, small, conical and slightly raised; a pair of white femoral glands, moderately large, rounded and slightly raised, at mid-femur.

Male with a pair of vocal slits on the posterior corners of the mouth, the single, external, vocal sac is indistinct on the preserved specimen; dark nuptial pads are present on the dorsomedial surface of the first and second fingers, from the base of the finger to the first terminal joint.

Colour. In life ( Fig. 3A–D ), the entire dorsal surface of the head, body, forelimbs and hindlimbs is light brown; a darker brown inverse triangular marking bounded by a thin fold is present from the interorbital region to the posterior area of the head. The dorsal surface of the body, forelimbs and hindlimbs have indistinct, darker brown blotches. The dorsal surface of the lower arms, tibia and femur (hindlimbs) have dark brown transverse folds. The lateral side of the head, including the canthal area, is brown, darker around the nostrils and just posterior and below the canthus, lighter anterior and posterior to the eye, with dark markings above and below the skin bordering the eye; supratympanic fold light brown, with indistinct dark brown blotches. The entire ventral surface is light brown with a heavy dark transverse marking and dark blotches over the throat and on the chest and abdomen. Ventral surface of throat to chest is dark brown, while the abdomen, forelimbs and hindlimbs are light cream, with some small blotches. In preservative ( Fig. 4A–E ) aspects of the colour pattern remain, but the dorsum darkens.

Variation (measurements in mm). Individuals of the type series are morphometrically generally similar ( Table 4 and 5). Based on the individual specimens collected, females appear to be slightly larger and stockier than males (male SVL 36.8–46.3 mm [n = 2] vs female SVL 38.2–79.1 mm [n = 4]). When the hindlimb is bent forward along the body, the tibiotarsal joint usually reaches to the posterior corner of the eye in adult males and reaches to the posterior of tympanum in females. Only one female (MZB Amph 3 472) had the tibiotarsal joint reach the posterior of the eye.

Individuals of the type series are only slightly variable in colouration. In preservative, the dorsal colouration in both males and females is relatively uniform light brown, but is completely dark brown in the single male paratype (MZB Amph 12564). The colouration of the ventral surface in both sexes is uniformly brown with dark blotches and dark longitudinal bands on the throat and chest. This longitudinal pattern is faded in one male (MZB Amph 12564).

Comparisons. Megophrys lancip sp. nov. differs from members of subgenera Xenophrys , Atympanophrys, Panophrys , and M. dringi by the absence of a folded gland forming a Y, X or H directed backwards at the parietoscapular region ( Günther, 1864; Delorme et al. 2006) (versus presence) ( Figs. 5A , 6C–D , Table 6).

Megophrys lancip sp. nov. is distinguished from all known members of Sundaland and Philippine Megophrys (except for M. montana and M. nasuta ), by the presence of a well -developed rostral appendage on the tip of the snout (versus absence in M. aceras , M. baluensis , M. dringi , M. edwardinae , M. longipes , M. kobayashii , and M. parallela ) ( Table 6, Fig. 5B–D ).

Megophrys lancip sp. nov. differs from Philippine M. stejnegeri by having a triangular-shaped eyelid appendage (versus a tubercle) and from M. ligayae by having a single dorsolateral fold that extends from the parietoscapular region to the groin ( Table 6, type I) (versus a dorsolateral folds that extends from the parietoscapular region to the middle of the body [ Table 6, type II]).

Megophrys lancip sp. nov. can be distinguished from M. nasuta by absence of additional lateral fold that extends from the area behind the supratympanic fold to the groin (versus presence in M. nasuta ) ( Table 6; Fig. 5D ), shorter rostral appendage SAL 1.5–4.1% of SVLh (versus SAL 1.2–9.2% of SVLh), and shorter eyelid appendage EHL 0.8–2.2 mm; 22.4% of UEWh (versus EHL 3.5–11.9 mm; 50.6% of UEWh) ( Table 5; Fig. 5B, D )

Megophrys lancip sp. nov. differs from M. montana by having a smaller body size, SVL 36.8–46.3 mm in adult males and 38.2–79.1 mm in adult females (versus 38.1–53.9 mm in adult males and 45.7–99.5 mm in adult females) ( Table 4), longer snout appendage, SAL 1.5–4.1% of SVLh, (versus SAL 0.0–3.5% of SVLh), shorter eyelid appendage EHL 0.8–2.2 mm; 22.4% of UEWh (versus EHL 0.9–3.9 mm; 30.0% of UEWh), shorter hand length, 8.6–10.4 mm in adult males and 9.8–19.0 mm in adult females, (versus 10.4–14.4 mm in adult males and 11.0– 23.6 mm in adult females), shorter hindlimb, HLL 52.0– 58.8 mm in adult males and 43.4–98.3 mm in adult females, (versus 56.2–76.6 mm in adult males and 56.0– 133.4 mm in adult females), tibio-tarsal articulation reached behind the eye in adult males and mostly reached behind the tympanum in adult females, (versus reached to the front of eye in adult males and reached the middle of eye in adult females), and less developed toe webbing I (1), II (1–1), III (2–2), IV (3–3), V (1 1/2) [versus I (1), II (1–1/2), III (1 3/4–1 1/2), IV (2 1/2–2 1/2), V(1)] ( Table 6).

TABLE ³. Variation in size anđ bođy proportions of male anđ female Megophrys from the Sunđalanđ anđ Philippines, mođifieđ from Taylor (1920), Inger (1954), Inger et al. (1995), Malkmus & Matsui (1997), anđ Inger & Iskanđar (2005).

* for M. parallela , the measurement đoes not incluđe the rostral appenđage.

……continued on the next page

……continued on the next page

……continued on the next page

Megophrys lancip sp nov. Megophrys montana

Species

RSAL

RHLh

RHW

RAG

REHL

RLAL

RHLL

RFML

RTL

REHL/UEWh

RTDH/ED

RFL/TL

RHW/HLh Species

male (n = 28) female (n = 21) juvenile (n = 12) male (n =6) female (n = 2)

RSAL 4.2 (1.5–8.7) 4.2 (1.8–9.2) 2.4 (1.2–5.4) – – – –

RHLh 44.0 (39.5–49.7) 46.0 (40.6–52.8) 42.0 (36.9–46.7) 42.6 (39.1–44.3) 41.8 (41.6–42.0) RHW 46.4 (44.0–49.5) 48.4 (44.8–54.8) 45.2 (37.0–49.4) 46.6 (44.9–51.9) 48.5 (48.5–48.6) RAG 42.0 (33.5–47.0) 46.5 (39.6–57.2) 48.4 (42.7–51.6) 44.5 (40.1–47.3) 46.3 (40.1–52.6) REHL 8.3 (6.3–11.4) 8.9 (4.4–13.0) 9.6 (6.0–12.7) 2.8 (1.8–3.4) 3.7 (3.2–4.2) RLAL 46.6 (20.8–49.8) 32.4 (19.9–48.2) 23.3 (20.7–25.2) 49.3 (46.3–50.5) 47.8 (47.2–48.5) RHLL 121.0 (108.8–147.9) 116.8 (106.6–126.2) 119.0 (111.6–129.0) 148.4 (139.9–161.5) 147.2 (143.0–151.4) RFML 40.5 (36.0–48.1) 38.9 (33.2–41.9) 40.4 (37.5–44.8) 48.1 (44.3–51.0) 47.5 (45.2–49.7) RTL 36.0 (33.3–44.1) 35.1 (31.8–37.9) 37.1 (34.4–40.4) 43.9 (41.3–48.4) 44.4 (42.4–46.4) REHL/UEWh 50 (38.5–61.4) 53.6 (32.7–61.3) 48.4 (32.1–62.7) 21.9 (14.7–29.0) 30.6 (24.9–36.3) RTDH/ED 32.7 (22.2–43.9) 35.7 (16.3–52.8) 28.2 (10.9–34.5) 32.4 (22.1–35.8) 31.5 (29.6–33.3) RFL/TL 111.8 (103.0–118.2) 111.1 (98.8–121.3) 108.9 (101–113.3) 107.1 (105.4–112.5) 106.9 (106.6–107.2) RHW/HLh 106.4 (92.9–116.1) 103.9 (92.4–119.5) 106.8 (91.6–111.2) 111.7 (106.9–118.1) 116.1 (115.4–116.9) Megophrys parallela Relative values (R) of each character compared to SVL (* used HL, and HW compared to HL).

* Fold forming Y, X or H on the parietoscapular region to the level of axilla.

** Dorsolateral fold shape:

Type I, dorsolateral folds are elongated and extend from the parietoscapular region to the groin.

Type II, dorsolateral folds extend from the parietoscapular region to the middle of the body

Type III, multiple dorsolateral folds - at least three or four - and they are discontinuous, formed by a series of elongated tubercles.

Type IV, dorsolateral folds are elongated and curve from the axillary region towards (and reaching) the posteriodorsal margin of tympanum.

Distribution and Natural History. The new species is known from the provinces of Lampung and Bengkulu in southwestern Sumatra ( Fig. 1 ). Larval, acoustic and other ecological data are unknown. The holotype was collected from a coffee plantation near the edge of secondary forest at 1010 m a.s.l. The male paratype from Bengkulu was collected from inside a rotten log on the forest floor at 620 m a.s.l. The female paratypes, collected from Bukit Barisan Selatan National Park, were collected from primary forest between 434 and 526 m a.s.l. This new species of Megophrys can be found sympatrically with M. nasuta in Kubu Prahu, Bukit Barisan Selatan National Park. Habitat loss and exploitation for the pet trade are likely be the main threats for the new species.

Figures

FIGURE 1. Map of southern Sumatra, showing known localities of the new species Megophrys lancip sp. nov..

FIGURE 2. Maximum Likelihood tree based upon a ~427 bp fragment of 16S rRNA gene for Megophrys lancip sp. nov., along with all representatives Sundaland and Philippine Megophrys species. Values at branches indicate Maximum Likelihood Bootstrap Proportion (MLBP), Bayesian Posterior Probabilities (BPP), and Neighbour Joining Bootstrap Proportion (NJBP).

FIGURE 3. Holotype of Megophrys lancip sp. nov. in life (adult male; MZB Amph 22233), showing (A) dorsolateral view, (B) chest, (C) ventral view and (D) dorsal view. Photos by E. N. Smith.

FIGURE 4. Megophrys lancip sp. nov. adult male holotype (MZB Amph 22233; A–E) and female paratype (MZB Amph 26738; F–J). Dorsal (A, F) and ventral (B, G) views of body, palmar views of right hand (C, H) and foot (D, I), and lateral view of head (E, J), scale bar = 5 mm.

FIGURE 5. Comparisons of male dorsal and ventral body views, left aspect of head, ventral aspects of head, hand and foot ventral views of: (A) M. aceras FMNH 268787 (photographs by Rachel Grill), (B) M. lancip sp. nov. UTA A-64763, (C) M. montana MZB Amph 6483, (D) M. nasuta UTA A-64764, and (E) M. parallela MZB Amph 13048 (formerly FMNH 264512), scale bar = 4 mm.

FIGURE 6. Comparisons of dorsal view of (A) M. montana (Syntype: RMNH.RENA 2212–A, (B) M. parallela (Paratype: MZB Amph 13048, formerly FMNH 264512), (C) M. monticola [reproduced from Günther (1864), plate XXVI figure H], and (D) M. parva [reproduced from Boulenger (1893), plate XI figure 2]. Red arrows added to show the parallel dorsolateral folds (A, B), or the Y-shaped folds from the parietoscapular region to the level of the axilla (C, D).