Limnonectes utara Matsui & Belabut & Ahmad 2014, sp. nov.

Limnonectes utara sp. nov. Synonymy: Rana kuhlii: Boulenger, 1912, p. 229 (part); Taylor, 1962, p. 408 (part); Berry, 1975, p. 71 (part). Rana kuhli: Inger, 1966, p. 196 (part). Limnonectes kuhlii: Grismer et al., 2010, p. 152. Holotype. UKMHC 528 (former KUHE 54064), an adult male from the upper part of Bukit Larut (= Larut Hill), State of Perak, Malaysia (4 o 51' N, 100 o 48' E, 1160 m a.s.l.: Fig. 10), collected on 28 February 2011 by Masafumi Matsui. Paratypes. KUHE 15447, 15463, 15465, 15514 (four adult females), KUHE 15442 (one young male), KUHE 15441 (one young female), and KUHE 15461 (one unsexed young) from lower part of Bukit Larut (680 m a.s.l.), collected on 3 January 1993 by Masafumi Matsui. KUHE 54065, 54089 (two adult males), KUHE 54086–54088 (three adult females), KUHE 54090 (one unsexed young) from the type locality, collected from 27 to 28 February 2011 by Masafumi Matsui and Norihiro Kuraishi. Referred specimens. UKMHC 700, 703, 704 from Sungai Tembat, Hulu Terengganu, State of Terengganu, Malaysia (ca. 5°26' N; 102°58' E, 168–170 m a.s.l.), collected in 2009 by Daicus M. Belabut; KUHE 49514 (three larvae) from lower part of Bukit Larut (600 m a.s.l.), collected on 3 January 1993 by Masafumi Matsui. Etymology. The specific epithet utara is a Malay word denoting north, alluding to the northerly-restricted distribution of the new species within the Peninsular Malaysia. Diagnosis. The new species is assigned to the genus Limnonectes only by molecular phylogenetic evidence, because morphological diagnostic characters of the genus previously proposed (e.g. Emerson and Berrigan 1993; Fei et al. 2005) are mostly osteological ones, which we did not examine. Moreover, osteological synapomorphic characters proposed by previous authors suffer from limited taxon sampling and require reexamination. Morphological assignment of the new species to Limnonectes was based only on the single characteristic usually found in the genus, enlarged “fangs” (tooth-like projections [odontoid processes]) on the lower jaw. A mediumsized species morphologically similar to L. kuhlii, with adult SVL 70–79 mm in males, 59–79 mm in females; males with relatively longer head than females; tibia dorsally densely covered by warts; toe webbing full, with very shallow excision between toes; first finger usually slightly longer than second, and nuptial pad present on first finger of males; morphologically differs from L. selatan, described below, in smoother dorsal skin, less densely arranged circum-cloacal warts, more confluent dorsal dark markings, and lack of large dark blotches on rear of thigh (Fig. 6). Description of holotype (measurements in mm). SVL 79.1; habitus stocky (Figs. 3A, 4A,B); head enlarged, longer (HL 36.0, 45.5%SVL) than broad (HW 33.8, 42.7%SVL); snout obtusely pointed, obtuse in profile, projecting beyond lower jaw; eye length (EL 11.0,13.9%SVL) shorter than snout length (SL 13.0, 16.4%SVL); canthus rounded; lore sloping, concave; nostril dorsolateral on canthus, closer to tip of snout than to eye; internarial distance (IND 5.8, 7.3%SVL) equal to upper eyelid (UEW 5.8, 7.3%SVL) and wider than interorbital distance (IOD 5.4, 6.8%SVL); pineal spot visible; tympanic annulus barely visible through skin; vomerine teeth in oblique groups, between and behind line connecting anterior rims of choanae, groups separated from one another by onefifth length of one group and from choana by about one-fourth length of one group; lower jaw with a pair of toothlike projections (odontoid processes) near symphysis, more than twice depth of mandible at base of projections; tongue oval, deeply notched posteriorly, without papillae; vocal sac and vocal slits absent. Forelimb heavy, relatively short (FLL 41.9, 53.0% SVL); fingers moderately slender; finger length formula: II <I <IV <III (Fig. 5A), first finger slightly longer than second, length of first (11.0, 13.9%SVL, measured from distal edge of inner palmar tubercle) equal to length of eye; fourth finger much longer than second; tips of fingers bluntly rounded, forming small pads without circummarginal grooves; no webs between fingers; inner palmar tubercle moderate (4.1, 5.2% SVL), oval, not elevated; middle palmar tubercle oval, smaller than inner palmar tubercle, not contacting outer or inner palmar tubercles; outer palmar tubercle slightly smaller than middle tubercle; proximal subarticular tubercles round and elevated; distal subarticular tubercles low and indistinct; no supernumerary metacarpal tubercles; narrow, but distinct flaps of skin along both edges of second and third fingers. Hindlimb heavy, relatively short (HLL 117.0,147.9% SVL) about 2.8 times length of forelimb; tibia short (TL 35.2, 44.5% SVL), heels not overlapping when limbs are held at right angles to body; tibiotarsal articulation of adpressed limb reaching to point posterior to eye; foot (FL 37.1, 46.9% SVL) slightly longer than tibia; toe length formula I <II <V <III <IV; tips of toes expanded into round, elevated pads lacking grooves (disk diameter of fourth toe, 4TDW 2.0, 2.5% SVL); all toes webbed to base of disks (Fig. 5B); webbing formula: I 0 – 0 II 0 – 0 III 0 – 0 IV 0 – 0 V; a distinct, movable flap of skin on outer edge of fifth toe and on inner edge of first toe; subarticular tubercles oval and distinct; an elongate inner metatarsal tubercle, length (IMTL 5.7, 7.2%SVL), about half length of first toe (1TOEL 11.2, 14.2% SVL); no outer metatarsal tubercle. Dorsum anteriorly very weakly rugose, without warts, posteriorly with very low ridges radiating from scattered, low warts (Fig. 4A); top of snout without tubercles, but eyelid covered with small warts and wrinkles with medial, longitudinal ridge formed by fused warts; weak transverse fold between posterior margins of eyes; a strong, supratympanic fold from eye to above axilla; posterior side of trunk scattered with tubercles; circum-cloacal warts small being sparsely and narrowly distributed anterior to cloaca, posteriorly more scattered but widely distributed, increasing size to end at dorso-ventral border of thigh (Fig. 6A); dorsal surface of thigh smooth on proximal two-thirds and scattered with small, low warts tipped with translucent spinules on distal one-third, continuing to tibia; tibia dorsally covered with numerous large and small round warts each tipped with large whitish cone surrounded by clusters of much smaller, whitish asperities; tarsus less densely covered by similar warts dorsolaterally; tarsus with a thick dermal ridge extending proximally from metatarsal tubercle; throat and chest weakly rugose, abdomen smooth (Fig. 4B); a distinct creamy tinge, with minute asperities, forming a nuptial pad covering medial surface of first finger from its base to level of subarticular tubercle, sharply set off from remainder of skin on first finger. Color. In life, dorsum light brown with confluent dark brown markings (Figs. 3A, 4A); head with a narrow light band anterior to dark interorbital bar; blackish brown stripe on canthus rostralis; side of head pale brown with dark markings; oblique blackish brown temporal stripe on and along supratympanic fold beginning behind eye reaching inguinal area, and continuing on flank; lips barred with dark brown; dark brown stripe on anterior side of upper arm; limbs marked dorsally with dark-brown crossbars; dorsal and ventral border of posterior thigh scattered with small brown spots (Fig. 6A); throat weakly mottled with pale gray, spotted with dark brown posteriorly; abdomen immaculate cream (Fig. 4B); ventral surfaces of hand and foot dark brown (Figs. 5 A,B). In preservative, the dorsal coloration has slightly faded, but otherwise no obvious change in color or pattern has occurred. Variation. Individual variation in size and body proportions is given in Table 3. Males have relatively longer head than females. Well-developed warts are invariably present, but there are some variations in qualitative traits. The postorbital light-colored bar was absent (50% of adults examined) or only vaguely traced (42%), and a few (8%) had a thin bar. The majority (83%) had a wide temporal stripe, and at least narrow temporal stripe was present (17%). Distinct spots were found widely distributed on dorsum in many (83%) individuals, and were found on at least part of the dorsum in others (17%). Many (75%) had weak spots or dots on the chin, but some had dusty marking (17%) or lacked chin spots (8%). The dark stripe on the upper arm was clear and continuous (38%) or weak or disjunct (46%), but was absent in some individuals (15%). Dorsal warts on the body were present usually (75%) only partially, and some (25 %) had very few warts. The first finger was usually longer than the second (69%), but was subequal (15%), or shorter than the first (15%) in some individuals. Eggs and tadpoles. The diameter of nine ovulated eggs from a female (KUHE 15447) ranged from 2.59–3.09 (mean±1SD = 2.74±0.17) mm. The animal hemisphere of the egg is dark brown and the vegetal hemisphere is pale yellow in color. Three tadpoles putatively assigned to the new species and at stage (Gosner, 1960) 35–36 (total length = 39.0–39.9 [39.6±0.5] mm, head-body length = 13.6–14.1 [13.8±0.3] mm) are nearly same as those of L. selatan described below in body shape and coloration (see Fig. 7). Comparisons. Limnonectes utara is most similar to L. selatan sp. nov., but is significantly larger in male SVL, and has relatively smaller internarial space, tibia, hindlimb, and first toe, and larger inner metatarsal tubercle and fourth toe disk. The dorsal dark markings are clearer and more confluent, marking on rear of thigh is less clear, and the circum-cloacal warts are less developed than in L. selatan (Fig. 6). The new species, along with L. selatan, is differentiated from all the other named species except for some Bornean L. kuhlii -like frogs by their tibia, which is heavily covered by large, conical tubercles (vs. tibia at most with weak tubercles dorsally). In addition, it differs from L. namiyei by the lack of vocal openings, and from L. fragilis by smaller subarticular tubercles and possession of nuptial pads in males. From L. kuhlii, it differs in possession of nuptial pads in males and larger body size (mean SVL 74 mm in males and 70 mm in females vs. 62 mm and 59 mm, respectively, in L. kuhlii). The new species, with a back usually without distinct ridges, and nuptial pad only on the first finger, differs from L. fujianensis, which has a back with many ridges, including a dorsolateral ridge, and nuptial pads on the two inner fingers. From L. bannaensis, it differs by lacking nuptial pad on the second finger. Limnonectes utara sp. nov. invariably has confluent dark dorsal marking unlike L. jarujini and L. isanensis, which usually lack dark dorsal markings. Limnonectes utara sp. nov. differs from L. taylori in having few dorsal warts, which are much more abundant and widely present in L. taylori. From L. megastomias, L. utara sp. nov. differs in having a smaller body size, relatively shorter head, lacking a nuptial pad on the second finger, and not having a heavily pigmented venter. In addition, the second finger is not constantly longer than first in the new species, unlike L. megastomias. Range. Peninsular Malaysia: Bukit Larut (= Larut Hill), State of Perak, (600–680 m a.s.l., 1160 m a.s.l.), Sungai Tembat, Hulu Terengganu, State of Terengganu, (168–170 m a.s.l.). Records of Rana or Limnonectes kuhlii from Larut Hills at about 4500 ft (Berry 1975), Ulu Kenas Recreational Forest, Gunung Bubu (Grismer et al. 2010), and Gunung Lawit (790 m), State of Terengganu (Dring 1979) are thought to represent this species (see Discussion). Known localities range in altitude from 168–1372 m a.s.l. Natural history. At the type locality (1160 m a.s.l.), the type series of L. utara sp. nov. was found in and along a small stream in a roadside ditch (width <1 m) in a secondary forest, while at the lower elevation site (680 m a.s.l.), frogs were found at the edge of the shaded main stream (width <3 m) in dipterocarp forest. Females collected in early January and late February had fully developed ova in ovaries. Tadpoles in later stages of development were also found in early January. Therefore, the breeding period may be relatively extended. Other amphibian species at the type locality were Megophrys longipes, Leptolalax heteropus, Ansonia malayana, Limnonectes khasianus, L. blythii, Amolops larutensis, Odorrana hosii, Hylarana banjarana, Philautus petersi, and Ichthyophis larutensis. Karyotype. The diploid chromosome number is 22, with six large and five small pairs (Matsui, unpub. data).Published as part of Matsui, Masafumi, Belabut, Daicus M. & Ahmad, Norhayati, 2014, Two new species of fanged frogs from Peninsular Malaysia (Anura: Dicroglossidae), pp. 75-93 in Zootaxa 3881 (1) on pages 80-86, DOI: 10.11646/zootaxa.3881.1.6, http://zenodo.org/record/494974

Limnonectes selatan Matsui & Belabut & Ahmad 2014, sp. nov.

Limnonectes selatan sp. nov. Synonymy: Rana kuhlii: Boulenger, 1912, p. 229 (part); Taylor, 1962, p. 408 (part); Berry, 1975, p. 71 (part). Rana kuhli: Inger, 1966, p. 196 (part). Holotype. UKMHC 529 (former KUHE 53961), an adult male from Genting Highlands, State of Pahang, Malaysia (3 o 24' N, 101 o 46' E, 1069 m a.s.l.: Fig. 10), collected on 22 February 2011 by Masafumi Matsui. Paratypes. KUHE 53379 (one adult female), KUHE 53380 (one young female) collected on 13 September 2009 by Daicus M. Belabut and Kanto Nishikawa, and KUHE 53962, 53963 (two adult males), KUHE 53964–53970, 53990 (eight adult females), KUHE 54010, 54011 (two unsexed young), KUHE 53955 (one just metamorphosed young) collected from 22 to 23 February 2011 by Masafumi Matsui, Daicus M. Belabut, Kanto Nishikawa, and Norihiro Kuraishi, all from the type locality. Referred specimens. KUHE 15106, 15108, 35567 (three unsexed young) from Ulu Gombak, State of Selangor, Malaysia (3 o 19' N, 101 o 45' E, ca. 200 m a.s.l.), collected on 9 December 1992 by Masafumi Matsui. KUHE 49515 (one lot of larvae, including KUHE 2011001 and 2011004 in Table 1), data same as the holotype. Etymology. The specific epithet selatan is a Malay word denoting south, alluding to the more southerlyrestricted distribution within the Peninsular Malaysia of the new species compared to L. utara. Diagnosis. Generic assignment of the new species is same as for L. utara described above. A medium-sized L. kuhlii -like species, with adult SVL 52–54 mm in males, 61–73 mm in females; males with relatively longer head than females; tibia densely covered by warts dorsally; toe webbing full, with very shallow excision between toes; first finger usually slightly longer than second, and nuptial pad present on first finger of males; differentiated from L. utara sp. nov. in having more rugose dorsal skin, more densely arranged circum-cloacal warts, dark dorsal spots usually isolated from other spots, and the presence of large dark blotches on the rear of the thigh. Description of holotype (measurements in mm). SVL 53.9; habitus stocky (Figs. 3B, 4C,D); head enlarged, slightly longer (HL 24.8, 46.0% SVL) than broad (HW 23.1, 42.9%); snout obtusely pointed, obtuse in profile, projecting beyond lower jaw; eye length (EL 7.8, 14.5% SVL) shorter than snout length (SL 8.8, 16.3% SVL); canthus rounded; lore sloping, concave; nostril dorsolateral, on canthus, closer to tip of snout than to eye; internarial distance (IND 5.1, 9.5% SVL) slightly smaller than interorbital distance (IOD 5.3, 9.8% SVL), latter wider than upper eyelid (UEW 4.4, 8.2% SVL); pineal spot visible; tympanic annulus slightly visible through skin; vomerine teeth in oblique groups, between and behind line connecting anterior rims of choanae, groups separated from one another by one-third length of one group and from choana by one-third length of one group; lower jaw with a pair of tooth-like projections near symphysis, more than twice depth of mandible at base of projections; tongue oval, deeply notched posteriorly, without papillae; vocal sac and vocal slits absent. Forelimb heavy, relatively short (FLL 28.5, 52.9% SVL); fingers moderately slender; finger length formula: II <I <IV <III, first finger slightly longer than second, length of first (6.7, 12.4% SVL, measured from distal edge of inner palmar tubercle) shorter than length of eye; fourth finger much longer than second; tips of fingers bluntly rounded, forming small pads without circummarginal grooves; no webs between fingers; inner palmar tubercle moderate (2.6, 4.8% SVL), oval, not elevated; middle palmar tubercle oval, smaller than inner palmar tubercle, not contacting outer or inner palmar tubercles; outer palmar tubercle slightly smaller than middle tubercle; proximal subarticular tubercles round and elevated; distal subarticular tubercles low, flat and indistinct; no supernumerary metacarpal tubercles; narrow, but distinct flaps of skin along both edges of second and third fingers. Hindlimb heavy, relatively short (HLL 80.5, 149.4% SVL) about 2.8 times length of forelimb; tibia short (TL 25.1, 46.6% SVL), heels not overlapping when limbs are held at right angles to body; tibiotarsal articulation of adpressed limb reaching to posterior corner of eye; foot (FL 26.5, 49.2% SVL) longer than tibia; toe length formula I <II <V <III <IV; tips of toes expanded into small, round, elevated pads lacking grooves (disk diameter of fourth toe, 4TDW 0.8, 1.5% SVL); all toes webbed to base of disks (Fig. 5D); webbing formula: I 0 – 0 II 0 – 0 III 0 – 0 IV 0 – 0 V; a distinct, movable flap of skin on outer edge of fifth toe and on inner edge of first toe; subarticular tubercles oval and distinct; an elongate inner metatarsal tubercle, length (IMTL 3.5, 6.5% SVL), less than half length of first toe (1TOEL 8.4, 15.6% SVL); no outer metatarsal tubercle. Dorsum anteriorly rugose, without warts, posteriorly with low ridges radiating from scattered, low warts; top of snout without tubercles; eyelid covered with small warts and wrinkles with medial, longitudinal ridge formed by fused warts; transverse fold between posterior margins of eyes absent; strong, supratympanic fold from eye to above axilla; side of trunk wrinkled and scattered with tubercles; circum-cloacal warts small, densely and widely distributed anteriorly, but increasing size posteriorly to end at dorso-ventral border of thigh (Fig. 6B); dorsal surface of thigh smooth on proximal two-thirds and scattered with small, low warts tipped with translucent spinules on distal one-third, continuing to tibia; tibia dorsally covered with numerous large and small round warts each tipped with a large whitish cone surrounded by clusters of much smaller, whitish asperities (Fig. 5C); tarsus less densely covered by similar warts dorsolaterally; tarsus with thick dermal ridge extending proximally from metatarsal tubercle; throat and chest weakly rugose, abdomen smooth; a distinct creamy tinge, with minute white asperities, forming a nuptial pad covering medial surface of first finger from its base to level of subarticular tubercle, sharply set off from remainder of skin on first finger. Color. In life, dorsum brown scattered with darker spots (Figs. 3B, 4C); head with a narrow light band anterior to dark interorbital bar; blackish brown stripe on canthus rostralis; side of head pale brown; oblique blackish brown temporal stripe on and along supratympanic fold beginning behind eye reaching inguinal area; lips barred with dark brown; dark brown spot on anterior side of upper arm; limbs marked dorsally with dark-brown crossbars; dorsal and ventral border of posterior thigh heavily marked with large dark brown spots (Fig. 6B); throat mottled with pale gray; abdomen immaculate cream (Fig. 4D); ventral surfaces of hand and foot dark brown (Fig. 5D). In preservative, the dorsal coloration has become darker, but otherwise no obvious change in color or pattern has occurred. Variation. Individual variation in size and body proportions is given in Table 3. In all individuals, postorbital light-colored bar was absent, and well-developed warts on the tibia and weak spots or dots on chin were present. Temporal stripe was usually narrow (in 92% of individuals), and was mere a trace in some (8%). Dorsal spots were usually few and weak (85%) and were absent in some (15%). A dark stripe on the upper arm was usually absent (85%), and was weak or disjunct in some (15%). Dorsal warts were usually present only partially (85%), and some had very few warts (15%). The first finger was usually longer than second (67%), but in some individuals they were subequal (17%), or the second was longer than first (17%). Eggs and tadpoles. Eggs and tadpoles from Genting Highlands were examined. The diameter of eight oviducal eggs from a female (KUHE 53379) ranged from 2.10–2.59 (mean±1SD = 2.37±0.18) mm. The animal hemisphere of egg is dark brown and the vegetal hemisphere is pale yellow in color. Each egg is enclosed with two transparent jelly layers. Tadpoles hatch at stage (Gosner 1960) 24 (total length = 10.7–12.2 [mean±1SD = 11.4±0.7] mm, head-body length = 3.7–3.8 [3.8±0.1] mm, n = 5) with rudimentary outer gills on left side, and can swim freely. They retain large amount of yolk and trace of suckers, and have a complete, but still unprotruded, eye under transparent skin and opened nostril, but the mouth is a mere depression and limb buds are absent. A total of five older tadpoles of stage 40 (total length=44.9–47.6 [mean±1SD = 46.4±1.2] mm, head-body length = 15.3–16.7 [16.1±0.5] mm) were closely examined. Head and body oval, slightly flattened above and below; width maximum at level of spiracle, 61–69 (median = 63)% of head-body length; depth 64–75 (median = 69)% of head-body width; snout rounded dorsally and in profile; eyes dorsolateral, not visible from below, eyeball diameter 11–12 (median = 12)% of head-body length; interorbital distance moderate, 116–136 (median = 136)% of eyeball diameter, less than eye-snout distance; nostril open, dorsolateral, rim not raised, midway between eye and tip of snout; internarial 69–83 (median = 74)% of interorbital. Oral disc anteroventral, emarginate, width 27–33 (median = 30)% of body width (Fig. 7H). Marginal papillae on upper labium with wide gap, lateral parts expanded, with eight short papillae in one row at corners, and few submarginal papillae; lower labium with continuous single row of wide, short and long papillae, and irregularly arranged submarginal papillae; labial teeth row formula 2(2)/ 3(1) or 2(2)/3(1–2), second anterior row rudimentary on each side, third posterior row about one-fourth length of second posterior row; serrated jaw sheaths forming wide arches with narrow black margins, upper jaw sheath with weak medial convexity. Spiracle sinistral, on left side, directed posterodorsally, tube fused to body wall, free portion distinct, length about half width of opening; snout-spiracle distance 43–57 (median = 54)% of head-body length. Vent tube dextral, attached to ventral fin. Tail lanceolate, both margins weakly convex, tapering gradually to tip; tail length 178–198 (median = 194)% of head-body length, maximum height 25–28 (median = 27)% of tail length; caudal muscle taller than either fin in proximal half of tail; origin of dorsal fin far posterior to end of body, dorsal fin higher than ventral fin except near tip of tail. Neuromasts of dorsal, infranaso-orbital, lateral, mental, pregular, postgular, postspiracle, and supranaso-orbital lines traceable. No glands present. In life, light brown dorsally with weak darker bands; dark band at junction of tail and body, continuing to base of caudal muscle. Fin and lateral caudal muscle speckled with large, dark transverse bands (Fig. 8). Two juveniles just after metamorphosis are 17.7 and 20.2 mm in SVL, with large conical warts on dorsal surface of tibia. Call characteristics. A call was recorded at the type locality at air and water temperature of 20.0°C and 19.8°C, respectively, at 18:50 h on 22 February 2011 by N. Kuraishi. However, the recording was poor quality because of heavy background noises of the stream, close to which the male stayed. The call consisted of a series of four notes and lasted 3.3 s. Each note was emitted at a gap (between the beginnings of two successive notes) of 0.94−1.24 (mean±1SD = 1.05±0.16) s and lasted for 0.12−0.14 (0.13 ± 0.01) s (Fig. 9). Pulse structure of each note could not be analyzed because of the poor quality of recording. The dominant frequency was ca. 0.5 kHz. Frequency modulation was observed within a note, and the frequency at the beginning 0.7–0.8 kHz decreased towards the end of the note to ca. 0.4 kHz. The frequency difference between the beginning and the end of the note was 0.3–0.4 kHz. Comparisons. Limnonectes selatan sp. nov. differs from the other L. kuhlii -like species in the same manner as L. utara sp. nov., by its uniquely developed tibial warts. It is very similar to L. utara sp. nov., but differs from it as noted above. Range. Peninsular Malaysia: Genting Highlands, State of Pahang (1069 m a.s.l.); Ulu Gombak, State of Selangor (ca. 200 m a.s.l.). Records of Rana kuhlii from Genting Simpah, Selanger (Berry 1975) and Hills of Selangor and Negri (sic.) Sembilan (Boulenger, 1912) are thought to represent this species (see Discussion). Known localities range in altitude from 200–1069 m a.s.l. Natural history: The type series of L. selatan was found at a swampy area in a valley, where there are seepages leading to a slowly moving small stream (width <2m) in the shade of shrubs and grasses in a well-grown secondary forest. In late February, occasional calls of males, females in ovulating condition, recently laid eggs, and larvae of various degree of development, including metamorphosed juveniles, were observed at the type locality. Among dead leaves on the bottom of shallow, slowly flowing water, eggs were laid scattered and tadpoles were found hiding themselves. Thus the breeding season seems last long at this locality unless the larval duration is long, which is unlikely. Amphibian species at the type locality were Megophrys longipes, Limnonectes blythii, L. tweediei, L. plicatellus, Odorrana hosii, Philautus petersi, and P. vermiculatus.Published as part of Matsui, Masafumi, Belabut, Daicus M. & Ahmad, Norhayati, 2014, Two new species of fanged frogs from Peninsular Malaysia (Anura: Dicroglossidae), pp. 75-93 in Zootaxa 3881 (1) on pages 86-89, DOI: 10.11646/zootaxa.3881.1.6, http://zenodo.org/record/494974

A New Species of Leptolalax (Amphibia, Anura, Megophryidae) from Peninsular Malaysia

A new megophryid species, Leptolalax kecil, is described from the Cameron Highlands of Peninsular Malaysia on the basis of acoustic and morphological characteristics. It has an advertisement call dissimilar to that of other, congeneric species, and is the smallest of the known Leptolalax. It is similar to L. pluvialis from Vietnam in small body size, but differs from it in ventral color, and in the size and color of the pectoral gland. The distributional pattern of the new species is discussed

Descriptions Of Two New Species Of Simulium (Simulium) (Diptera: Simuliidae) From Tioman Island, Peninsular Malaysia

Takaoka, Hiroyuki, Sofian-Azirun, Mohd, Belabut, Daicus M. (2012): Descriptions Of Two New Species Of Simulium (Simulium) (Diptera: Simuliidae) From Tioman Island, Peninsular Malaysia. Raffles Bulletin of Zoology 60 (2): 399-409, DOI: 10.5281/zenodo.534971

Variation In The Nucleolar Organiser Regions Of The Long-Tailed Giant Rats (Rodentia, Muridae, Genus Leopoldamys) In Malaysia

Yong, Hoi Sen, Lim, Phaik-Eem, Belabut, Daicus M., Eamsobhana, Praphathip (2013): Variation In The Nucleolar Organiser Regions Of The Long-Tailed Giant Rats (Rodentia, Muridae, Genus Leopoldamys) In Malaysia. Raffles Bulletin of Zoology 61 (2): 855-859, DOI: http://doi.org/10.5281/zenodo.535312

Morphological Divergence, Reproductive Isolating Mechanism, and Molecular Phylogenetic Relationships Among Indonesia, Malaysia, and Japan Populations of the Fejervarya limnocharis Complex (Anura, Ranidae)

In order to elucidate the taxonomic status of the Fejervarya limnocharis complex relative to Malaysia and Japan populations, morphological observations and molecular phylogenetic analysis were carried out using three populations from Indonesia (type locality), Malaysia, and Japan. In addition, we conducted histological and spermatogenic observations using hybrids among these populations. Principal component and cluster analyses demonstrated that these populations could be clearly separated from one another. Abnormal testes were found in the hybrids between the Japan and Indonesia populations and between the Japan and Malaysia populations, but testes of the controls and hybrids between the Malaysia and Indonesia populations were quite normal. The mean number of univalents per cell was 5.42, 4.58, and 0.20 in hybrids between the Indonesia and Japan populations, Malaysia and Japan populations, and Indonesia and Malaysia populations, respectively. Sequence divergences in 16S rRNA and Cyt b genes were 0–0.4% (x–=0.2%) and 0.3–1.5% (x–=1.0%), respectively, between the Malaysia and Indonesia populations, and 2.4–2.6% (x–=2.5%) and 11.0–12.0% (x–=11.5%) between the Japan population and F. limnocharis complex, including the Malaysia and Indonesia populations and F. multistriata from China. This study indicated that the Malaysia population and F. multistriata from China should be designated as a subspecies of topotypic F. limnocharis, and that the Japan population should be regarded as a distinct species

Morphological Divergence, Reproductive Isolating Mechanism, and Molecular Phylogenetic Relationships Among Indonesia, Malaysia, and Japan Populations of the Fejervarya limnocharis Complex (Anura, Ranidae)

In order to elucidate the taxonomic status of the Fejervarya limnocharis complex relative to Malaysia and Japan populations, morphological observations and molecular phylogenetic analysis were carried out using three populations from Indonesia (type locality), Malaysia, and Japan. In addition, we conducted histological and spermatogenic observations using hybrids among these populations. Principal component and cluster analyses demonstrated that these populations could be clearly separated from one another. Abnormal testes were found in the hybrids between the Japan and Indonesia populations and between the Japan and Malaysia populations, but testes of the controls and hybrids between the Malaysia and Indonesia populations were quite normal. The mean number of univalents per cell was 5.42, 4.58, and 0.20 in hybrids between the Indonesia and Japan populations, Malaysia and Japan populations, and Indonesia and Malaysia populations, respectively. Sequence divergences in 16S rRNA and Cyt b genes were 0–0.4% (x̄=0.2%) and 0.3–1.5% (x̄=1.0%), respectively, between the Malaysia and Indonesia populations, and 2.4–2.6% (x̄=2.5%) and 11.0–12.0% (x̄=11.5%) between the Japan population and F. limnocharis complex, including the Malaysia and Indonesia populations and F. multistriata from China. This study indicated that the Malaysia population and F. multistriata from China should be designated as a subspecies of topotypic F. limnocharis, and that the Japan population should be regarded as a distinct species

From Antarctica or Asia? New colonization scenario for Australian-New Guinean narrow mouth toads suggested from the findings on a mysterious genus Gastrophrynoides

Background: Microhylidae is a geographically widespread family of anurans. Although several extensive molecular analyses have attempted to elucidate their subfamilial relationships, and correlate these with Mesozoic and Cenozoic continental drifts, consensus has not been reached. Further, generic level relationships have not been well investigated in some microhylid subfamilies, and therefore subfamilial affiliations of some genera are still unclear. To elucidate the phylogenetic positions of two mysterious Asian genera, Gastrophrynoides andPhrynella, and to better understand the trans-continental distributions of microhylid taxa, we performed molecular phylogenetic and dating analyses using the largest molecular dataset applied to these taxa to date. Results: Six nuclear and two mitochondrial genes (approx. 8 kbp) were sequenced from 22 microhylid frog species representing eight subfamilies. The maximum likelihood and Bayesian analyses could not fully elucidate the subfamilial relationships, suggesting a rapid radiation of these taxa between 85 and 66 million years ago. In contrast, generic relationships of Asian microhylines were generally well resolved. Conclusion: Our results clearly showed that one of two problematic Asian genera, Phrynella, was nested in the clade of the Asian subfamily Microhylinae. By contrast, Gastrophrynoides occupied the most basal position of the Australian-New Guinean subfamily Asterophryinae. The estimated divergence of Gastrophrynoides from other asterophryine was unexpectedly around 48 million years ago. Although a colonization scenario via Antarctica to the Australian-New Guinean landmass has been suggested for Asterophryinae, our finding suggested a novel colonization route via Indo-Eurasia