6,186 research outputs found
Meristogenys amoropalamus Matsui 1986
Meristogenys amoropalamus (Matsui, 1986) Amolops amoropalamus: Matsui, 1986, p. 628–629. figs. 1 C, 1 F, 2 C, 2 D; Meristogenys amoropalamus (larvae): Yang, 1991, p. 33–34; Malkmus et al., 2002, p. 149–150; Meristogenys sp. (lineage 1): Shimada et al., 2007 a, pp. 173–189; Meristogenys amoropalamus (larval morphotype 1): Shimada et al., 2007 b, pp. 59–63; Meristogenys amoropalamus (lineage 1): Shimada et al., 2011, p. 158–177, Fig. 3, 4, 6. Iris colour. Iris bicoloured, yellowish green above and reddish brown below. The upper part became dark yellow after anaesthesia. Eggs. Ova bicoloured, heavy pigmentations on the animal pole, cream white on the plant pole; clutch size 400– 550; ovum diameter 2.1–2.6 mm. Range. Besides the type locality (Gunung Tapai Sia; 1300 m a. s. l.), the adult of this species has been collected from Mesilau (1800 m) and Kamborangah (1800 m) in the Kinabalu National Park and Bario (1000 m) from Sarawak. The larvae of this species have also been collected from Mahua (1063 m), Liwagu (1500 m), and Trus Madi (Shimada et al., 2007 b; altitude unknown). According to Yang (1991), tadpoles similar to this species had been collected from Sg. Panataran [Melangkap; FMNH (Field Museum of Natural History) 229861], Sg. Mesilau (Mesilau; FMNH 229862), Sg. Bambangan (Mamut; FMNH 229863), Sg. Kipungit (Poring; FMNH 130893), and Sg. Pegalan (FMNH 228007; J. Ladonski, pers. comm.).Published as part of Shimada, Tomohiko, Matsui, Masafumi, Yambun, Paul & Sudin, Ahmad, 2011, A survey of morphological variation in adult Meristogenys amoropalamus (Amphibia, Anura, Ranidae), with a description of a new cryptic species, pp. 33-56 in Zootaxa 2905 on page 51, DOI: 10.5281/zenodo.20559
Congressman Bob Matsui
Scan of photo signed "To Mike, best wishes to my friend, Bob Matsui." Matsui, from Sacramento, California, served as a Congressman from 1979 to 200
77. Matsui Sumako (1886-1919)
Iwao Seiichi, Iyanaga Teizō, Ishii Susumu, Yoshida Shōichirō, Fujimura Jun'ichirō, Fujimura Michio, Yoshikawa Itsuji, Akiyama Terukazu, Iyanaga Shōkichi, Matsubara Hideichi. 77. Matsui Sumako (1886-1919). In: Dictionnaire historique du Japon, volume 14, 1988. Lettres L et M (1) p. 45
77. Matsui Sumako (1886-1919)
Iwao Seiichi, Iyanaga Teizō, Ishii Susumu, Yoshida Shōichirō, Fujimura Jun'ichirō, Fujimura Michio, Yoshikawa Itsuji, Akiyama Terukazu, Iyanaga Shōkichi, Matsubara Hideichi. 77. Matsui Sumako (1886-1919). In: Dictionnaire historique du Japon, volume 14, 1988. Lettres L et M (1) p. 45
Leptolalax fritinniens Dehling & Matsui, 2013, sp. n.
Leptolalax fritinniens sp. n. Twittering Litter Frog (Figures 1–4) Holotype: NMBE 1056267, adult male, from near Camp 5 (4 °08.208' N, 114 ° 53.622 ' E, ca. 114 m a.s.l.), Gunung Mulu National Park, Miri Division, Sarawak, East Malaysia (Borneo), collected 3 December 2007 by J. M. Dehling. Paratypes: BMNH 1978.1524, 1978.1525, two adult females, from Batu Pala, a limestone outlier near Camp 5, Gunung Mulu National Park, collected in 1977 by Julian M. C. Dring; NMBE 1056367–1056368, two adult males, from near Camp 5, Gunung Mulu National Park, collected 20 March 2009 by A. Haas, J. M. Dehling, Pui Y.-M., S. T. Hertwig, and A. Jankowski; KUHE 10534, an adult male, from Camp 5, Gunung Mulu National Park, collected 26 December 1989 by M. Matsui and K. Araya; SRC unnumbered (former KUHE 53678), KUHE 53676, 53679– 53681, four adult males, from near Camp 5, Gunung Mulu National Park, collected 21 August 2010 by M. Matsui, K. Nishikawa, and K. Eto. Diagnosis. We allocate the new species to the genus Leptolalax for showing the following diagnostic characters: vomerine teeth absent, finger tips rounded, fingers lacking webbing, with toe webbing basal or rudimentary, limbs relatively long and slender, subarticular tubercles indistinct, inner palmar tubercle elevated and not extending to Finger I, outer metatarsal tubercle absent, nuptial pads absent, tip of snout with vertical white bar (Dubois 1983, 1987; Lathrop et al. 1998, Delorme et al. 2006). The new species can be distinguished from all other members of the genus Leptolalax by the combination of following characters: (1) size large, SVL of males 31.8 – 34.0 mm, of females 45.5–47.7 mm; (2) snout rounded in both ventral view and lateral view; (3) interorbital distance smaller than width of upper eyelid; (4) vocal sac of males subgular, bipartite; (5) toe webbing basal; (6) skin on dorsum and dorsal side of head shagreened with tiny tubercles; (7) supratympanic fold angled; (8) pectoral glands small; (9) supraaxillary glands and ventrolateral glandular ridges absent; (10) venter spotted; (11) advertisement call consisting of long series of 8–289 notes, each composed of with three or four pulses, dominant frequency at 7225–9190 Hz, with prominent frequency modulation. Etymology. The species epithet derives from the Latin fritinniens, meaning “twittering”, in allusion to the species’ high-pitched advertisement call with prominent frequency modulation. Description of holotype. Habitus moderately slender (Figures 1 & 2); head moderately wide (HW/SVL 0.32), about as wide as long (HW/HL 0.98) and wider than trunk; snout rounded in both ventral view and lateral view, slightly protruding, its length 40 % of head length and 90 % of eye diameter, wider than long (SL/EE 0.84); canthus rostralis distinct, slightly concave between eye and nostril in dorsal view, almost straight-lined in lateral view; loreal region oblique, slightly concave; nostrils oval, directed dorsolaterally, closer to tip of snout than to eye (EN/ NS 1.12); distance between eye and nostril subequal to internarial distance (EN/NN 0.97) and much smaller than eye diameter (EN/ED 0.58); eye very large (ED/HL 0.45); pupil vertical; tympanum distinct, rounded, its diameter half the eye diameter (TD/ED 0.50); interorbital distance smaller than width of upper eyelid (IO/EW 0.65) and smaller than internarial distance (IO/NN 0.90); pineal ocellus absent; symphysial knob on anteriormost part of mandible; vomerine ridge and teeth absent; tongue large, broad, bifid, free for about half its length; median lingual process absent; vocal sac subgular, bipartite, consisting of two lateroventrally situated parts which are fused with each other medially when inflated (Figure 3); apertures of vocal sac slit-like, directed posterolaterally, situated halfway between base of tongue and corners of mouth. Forelimbs slender, moderately long (ELB/SVL 0.70, ARM/SVL 0.56); hand longer than forearm (HND/ARM 0.55); fingers long and slender, without webbing or lateral fringes of skin (Figure 4); relative length of fingers II<I<IV<III; finger tips rounded and thickened; subarticular tubercles indistinct; large, prominent, rounded tubercle in thenar and metacarpal region of fingers I, II, and III, separated by a distinct groove from much smaller, rounded tubercle in metacarpal region of Finger IV. Hindlimbs moderately long (LEG/SVL 1.69); tibiofibula long (TFL/SVL 0.54), longer than foot (TFL/FOT 1.12) and longer than thigh (TFL/THL 1.07); heels overlapping each other with knees flexed and thighs being held perpendicularly to median plane of body; toe tips rounded and thickened, smaller than finger tips; toe webbing basal, webbing formula I 2 -2.5 II 2- 3 III 2.75- 4 IV 4 +- 3 V (Figure 4); narrow fringes of skin on lateral sides of toes; relative length of toes I<II<V<III<IV; subarticular tubercles indistinct; longitudinal ridges of thickened skin on plantar side of phalanges except distal ones of toes II–V, absent on Toe I; inner metatarsal tubercle large (length 2.0 mm), oval, 44 % of length of first toe; outer metatarsal tubercle absent. Skin on dorsum and dorsal side of head shagreened with tiny tubercles, weakly wrinkled on dorsal surfaces of the extremities (Figures 1, 2); wrinkles on extremities forming indistinct, reticulated, predominantly longitudinal, low ridges (Figures 1, 2, 3); several enlarged tubercles on lateral surfaces of trunk; ventral side smooth; supratympanic fold thick and conspicuous, angled, running from posterior margin of eye to just behind corner of mouth; pectoral glands small, indistinct, at insertion of forelimbs; supraaxillary glands and ventrolateral glandular ridges absent. Measurements. SVL 33.1, TFL 17.8, FOT 15.8, TarL 24.9, LEG 55.9, THL 16.5, ELB 23.2, ARM 18.4, HND 10.1, HW 10.7, HL 10.9, IO 2.6, EW 4.0, ED 4.9, TD 2.4, TE 1.1, EN 2.8, NS 2.5, SL 4.4, NN 2.9, EE 5.2. Colouration in life. Basic colouration of dorsum anthracite with several light brown flecks and large black, irregularly shaped spots; area below canthus rostralis and ventral edge of supratympanic fold black; basic colouration lightened to light grey on lateral surfaces of trunk and almost white on venter; dorsal surfaces of extremities light grey with dark grey, black-edged crossbars; dorsal area at tibio-tarsal articulation cream-coloured; several cream-coloured stripes along edges of jaws and similarly coloured crossbars on upper arm and on dorsal surface of fingers and toes; conspicuous, light grey stripe from between nostrils to anteriormost edge of upper jaw; venter with large light grey spots; throat, anterior portion of breast, ventral side of forelimb moderately densely speckled, ventral surface of thigh, tibia, tarsus, and postaxial sides of forelimbs heavily speckled dark brown; posterolateral portion of the throat where vocal sacs are located largely unpigmented; iris dark grey in ventral twothirds, reddish grey in dorsal third, with inner edge around pupil ruby-red. Colouration in preservative. Colours generally darker; contrast reduced; pattern still distinct; iris colour faded to bluish-grey. Variation. The male paratypes are generally very similar to the holotype in size and proportions. SVL of males varies between 31.8 mm (NMBE 1056367) and 34.0 mm (NMBE 1056368). Females are considerably larger than males with an SVL of 45.5 mm (BMNH 1978.1524) and 47.7 mm (BMNH 1978.1525). Males have vocal sacs but lack nuptial pads or asperities. TFL/SVL ratio is 0.53–0.56 (males) and 0.47–0.50 (females), HW/HL 0.96–1.03 (males) and 1.13–1.14 (females), IO/EW 0.65–0.87 (males) and 0.70–0.75 (females), and EN/NN 0.97–1.08 (males) and 1.05–1.06 (females). Advertisement call. Advertisement calls of four males including the holotype were recorded and analysed (Table 1). Values are given as mean ± SD followed by range. Air temperatures during recordings ranged between 24.3 and 24.9 °C. The advertisement consisted of long series of notes (Figure 5). Number of notes within a series was 58.7 ± 36.3 (8–289). Note repetition rate was 11.7 ± 0.3 (10.6–12.8) per second. Depending on the number of notes, calls lasted 4.9 ± 2.5 (0.6–17.4) s. Individual notes lasted 34.0 ± 4.2 (14–50) ms and were separated from each other by an interval of 51.3 ± 4.7 (36–61) ms. Each note was composed of 3.3 ± 0.5 (3–4, n = 30) rather indistinct pulses (Figure 5 C). Pulse length varied considerably between 3 and 17 ms (8.8 ± 3.7, n = 37). When pulses were discrete, the interval between individual pulses was about 2 ms. Marked frequency modulation was always observed within a note. The dominant frequency at the beginning was 8266 ± 250 (7750–9200) Hz and decreased towards the end of the note to 7622 ± 154 (7250–8100) Hz. The frequency difference between the beginning and the end of the note was 650 ± 150 (200–1200) Hz. Usually, the first note or the first two notes of a series were higher in frequency (9142 ± 90 Hz, n = 129 calls from 2 males) than the subsequent ones (Figure 5 B, C). Irregular intensity modulation was observed within a single call and a note (Figure 5). Only in the recording of one male, weak harmonics were present between 9500–9800 Hz. In phases of few calls of this male, only these harmonics were audible and the corresponding notes were only about half the length of regular notes. During the recording, this male sometimes continued the vocal sac motions between two calls but no sound was audible or traceable between 20 and 44,000 Hz in the recordings. Ecological notes and distribution. The type specimens were found in alluvial forest perching on leaves in low vegetation in the vicinity of small, slow-flowing streams. All males were encountered while calling. Most specimens were found at night, but a few were heard calling between 0 930 h and 1130 h. Tadpoles of the species remain unknown. The species occurs sympatrically with Leptolalax pictus and Leptolalax cf. gracilis. Although the type series contains only specimen from Camp 5 in Gunung Mulu National Park, we are aware that the species is also distributed in Brunei and Sabah (unpublished molecular and bioacoustic data of MM and JMD). However, most voucher specimens in herpetological collections are mislabelled as either L. dringi or L. gracilis. In addition, preliminary genetic and bioacoustic analyses of specimens and their calls from several locations in Sabah (Matsui, unpubl. data; Dehling, unpubl. data) indicate that at least one further, yet unrecognized species is present. Before the relationships between the populations from Sabah have been resolved, we therefore refrain from describing the geographic range of the new species. (0.6–15.4, n = 111) (1.1–7.2, n = 11) (1.8–3.5, n = 5) (0.9–17.4, n = 18) call interval [s] 2.7 ± 2.1 1.3 ± 0.5 8.7 ± 4.4 1.6 ± 0.4 (1.3–19.7, n = 111) (0.4–1.9, n = 10) (5.0– 14.5, n = 4) (1.2–2.5, n = 16) number of notes/call 49.0 ± 32.0 55.8 ± 24.1 28.2 ± 7.3 98.9 ± 89.4 (8–186, n = 111) (14–87, n = 11) (21–38, n = 5) (12–289, n = 18) note repetition rate [/s] 11.8 ± 0.2 11.9 ± 0.5 11.3 ± 0.4 11.8 ± 0.2 (11.2–12.1, n = 111) (10.62–12.79, n = 11) (10.87–11.76, n = 5) (11.3–12.1, n = 18) note duration [ms] 34.7 ± 4.5 35.0 ± 3.2 38.0 ± 5.8 28.1 ± 3.4 (28–44, n = 90) (29–44, n = 30) (24–50, n = 31) (14–35, n = 90) note interval [ms] 47.9 ± 4.8 47.0 ± 3.8 54.6 ± 5.2 54.6 ± 3.8 (550–750, n = 90) (200–1200, n = 30) (400–950, n = 31) (500–1100, n = 90) Comparisons. By the absence of both a ventrolateral glandular ridge and a supraaxillary macrogland, Leptolalax fritinniens differs from species of the genus occurring north of the Isthmus of Kra (in most parts corresponding to the subgenus Lalos Dubois, Grosjean, Ohler, Adler & Zhao, 2010), all of which have a ventrolateral glandular ridge and/or a supraaxillary gland, i.e. Leptolalax aereus Rowley, Stuart, Richards, Phimmachak & Sivongxay, 2010, L. alpinus Fei, Ye & Li, 1990, L. applebyi Rowley & Cao, 2009, L. bidoupensis Rowley, Le, Tran & Hoang, 2011, L. bourreti Dubois, 1983, L. croceus Rowley, Hoang, Le, Dau & Cao, 2010, L. eos Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011, L. firthi Rowley, Hoang, Dau, Le & Cao, 2012, L. fuliginosus Matsui, 2006, L. khasiorum Das, Tron, Rangad & Hooroo, 2010, L. lateralis (Anderson, 1871), L. liui Fei & Ye, 1990, L. melanoleucus Matsui, 2006, L. melicus Rowley, Stuart, Thy & Emmett, 2010, L. nahangensis Lathrop, Murphy, Orlov & Ho, 1998, L. nyx Ohler, Wollenberg, Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011, L. oshanensis (Liu, 1950), L. pelodytoides (Boulenger, 1893), L. pluvialis Ohler, Marquis, Swan & Grosjean, 2000, L. sungi Lathrop, Murphy, Orlov & Ho, 1998, L. tamdil Sengupta, Sailo, Lalremsanga, Das & Das, 2010, L. tuberosus Inger, Orlov & Darevsky, 1999, and L. ventripunctatus Fei, Ye & Li, 1990. From the species occurring on the Malay Peninsular and Borneo, L. fritinniens (characters in parentheses) differs in the following morphological characters: Leptolalax arayai Matsui, 1997 has a tuberculate dorsum (vs. smooth) without conspicuous markings (vs. present in L. fritinniens), has an unspotted, yellow venter (vs. venter white with black spots), orange groin and ventral sides of legs (vs. white to cream-coloured ventral side of legs and groin), and a single, medially arranged vocal sac in males (vs. bipartite; Figure 3). Leptolalax dringi Dubois, 1987 differs by a smaller size with SVL 26.6–31.3 mm in males and 36.6–38.1 mm in females (vs. SVL 31.8 –34.0 mm in males and 45.5–47.7 mm in females); iris bright red in upper third (vs. reddish grey); a relatively wider interorbital space with IO/EW 0.80–1.04 in males and 0.74–0.79 in females (vs. 0.65–0.87 and 0.70–0.75, respectively); angle of supratympanic fold being wider (vs. angle narrow); heads of males being wider than long with HW/HL 1.11–1.22 (vs. as long as wide HW/HL 0.96–1.03); and a relatively greater eye-to-nostril distance with EN/NN 0.81–0.90 in females and 0.76–0.89 in males (vs. 1.05–1.06 in females and 0.97–1.08 in males). Leptolalax gracilis (Günther, 1872) differs in having a curved supratympanic fold (vs. angular); a single medially arranged subgular vocal sac in males (vs. bipartite; Figure 3); relatively greater interorbital distance with IO/EW 0.78–0.99 in females and 0.71–0.89 in males (vs. 0.70–0.75 and 0.65–0.87, respectively); iris bright red in the upper two-fifths and dull greyish red, with a narrow ring of bright red along the pupil in the lower three-fifths (vs. iris dark-grey in ventral two-thirds, reddish grey in dorsal third, with inner edge around pupil ruby-red); male SVL 34.3 –39.0 (vs. 31.8 –34.0). Leptolalax hamidi Matsui, 1997 differs in having the venter unspotted (vs. spotted); large dark brown dorsal markings with light outlines (vs. dorsal markings small without light outlines); and a smaller size with SVL 28.0–31.0 mm in males, 36.0–43.0 mm in females (vs. 31.8 –34.0 mm in males and 45.5– 47.7 mm in females). Leptolalax heteropus (Boulenger, 1900) is much smaller with SVL of males 24.6 –26.0 mm and of female 31.7 mm (vs. 31.8 –34.0 mm in males and 45.5–47.7 mm in females) and has more developed toe webbing and distinct lateral fringes on the toes (vs. toe webbing basal, lateral fringes indistinct). Leptolalax kajangensis Grismer, Grismer & Youmans, 2004 has shorter legs with TFL/SVL 0.42 in males (vs. 0.53–0.56); a curved supratympanic fold (vs. angular); a black spot on the tympanum (vs. absent); an unpatterned venter (vs. venter spotted); and a dark brown dorsum with black pattern (vs. grey dorsum). Leptolalax kecil Matsui, Belabut, Ahmad & Yong, 2009 is much smaller with SVL of males 19.3–20.5 mm and female 25.0 mm (vs. 31.8 –34.0 mm in males and 45.5–47.7 mm in females), has a light brown dorsum with dark brown pattern (vs. grey dorsum with black spots); a black spot on the tympanum (vs. absent); a red iris (vs. iris dark grey in ventral two-thirds, reddish grey in dorsal third); and has large, conspicuous, orange pectoral glands (vs. pectoral glands hardly discernible). Leptolalax maurus Inger, Lakim, Biun & Yambun, 1997 is much smaller with SVL of the female holotype being 31.8 mm (vs. 43.5–47.7 mm in females) and SVL of the male paratype 26.1 mm (vs. 31.8 –34.0 mm in males), has a dark brown to black dorsal and ventral colouration (vs. grey dorsally and white with black spots ventrally); and a dark red iris (vs. dark grey in ventral two-thirds, reddish grey in dorsal third). Leptolalax pictus Malkmus, 1992 has an immaculate venter (vs. spotted) and has a light brown dorsum with dark brown markings with conspicuous thin light outlines (vs. dorsum grey with black pattern without distinct light outlines). Leptolalax platycephalus Dehling, 2012 has an immaculate venter (vs. spotted); a skin flap above the vent; a wider head with HW/SVL 0.37–0.38 (vs. 0.31–0.32); a greater interorbital distance with IO/EW 1.14–1.27 (vs. 0.65–0.87), rudimentary toe webbing (vs. basal), large pectoral glands (vs. hardly discernible), and different dorsal and ventral colouration. Leptolalax solus Matsui, 2006 is smaller with the only specimen, a male, having an SVL of 27.6 mm (vs. 31.8 –34.0 mm in males); has a larger pectoral gland; and a chocolate-brown, largely unpatterned dorsum (vs. dorsum grey with dark pattern). In addition to the morphological differences, Leptolalax fritinniens can be distinguished by its unique advertisement call from 20 of the 35 species of the genus, of which call characteristics are known (Malkmus & Riede 1993; Matsui 1997; Jiang et al. 2002; Malkmus et al. 2002; Xu et al. 2005; Matsui 2006; Matsui et al. 2009; Rowley & Cao 2009; Sukumaran et al. 2010; Rowley et al. 2010 a, 2010 b, 2010 c, 2011, 2012; Matsui & Dehling 2012). The advertisement call of L. fritinniens, as described by Matsui (1997), was compared to calls of recently described species (as L. dringi; Matsui 1997, 2006; Matsui et al. 2009; Rowley & Cao 2009; Rowley et al. 2010 a, 2010 b, 2010 c, 2011, 2012), and therefore, we compare the advertisement call only to those of the other Bornean species in the following: The call of L. fritinniens has a dominant frequency of 7250–9200 Hz at 24.3–24.9 °C. Although dominant frequency is known to vary over temperature (e.g. Rowley et al. 2010 c) the frequency of the call of L. fritinniens is likely to be higher than the dominant frequency of the calls of L. arayai (5400–5900 Hz, 17.4 °C), L. dringi (6050–6400 Hz, temperature unknown), L. gracilis (2600–2800 Hz, 20.0– 26.2 °C), L. hamidi (6700–7300 Hz, 22.9–24.1 °C), L. maurus (5150 Hz, temperature unknown), and L. pictus (6800–7150 Hz, 19–22 °C). Call length and number of notes within a call is very variable (0.6– 17.4 s, 8–289 notes), but on average (4.9 s, 59 notes), the calls recorded in L. fritinniens are longer and contain more notes than reported in the call of L. dringi (0.1 – 1.0 s, 2–11 notes), L. maurus (4.1 s, 15 notes), and L. pictus (1.9– 4.9 s, 12–31 notes). Marked frequency modulation like in the calls of L. fritinniens is absent in the calls of L. dringi, L. gracilis, and L. maurus. Note repetition rate is temperature-dependent but at 10.6–12.8 per second recorded for L. fritinniens at 24.3–24.9 °C it appears to be higher than in the advertisement calls of all other Bornean species (L. arayai 9.0– 9.3 /s; L. dringi 8.2– 10.3; L. gracilis 6.9–10.4; L. hamidi 9.0– 9.3; L. maurus 3.5) except L. pictus (11–13).Published as part of Dehling, J. Maximilian & Matsui, Masafumi, 2013, A new species of Leptolalax (Anura: Megophryidae) from Gunung Mulu National Park, Sarawak, East Malaysia (Borneo), pp. 33-44 in Zootaxa 3670 (1) on pages 35-41, DOI: 10.11646/zootaxa.3670.1.2, http://zenodo.org/record/28392
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
Meristogenys dyscritus Shimada, Matsui, Yambun & Sudin, 2011, sp. nov.
Meristogenys dyscritus sp. nov. Amolops sp. A: Matsui, 1979, p. 340, figs. 26, 28A; Amolops amoropalamus: Matsui, 1986, p. 628–629 (part); Meristogenys amoropalamus (lineage 3, lineage 4): Shimada et al., 2007 a, p. 173–189; Shimada et al., 2008, p. 24–34; Shimada et al., 2011, p. 158–177, Fig. 3–4, Appendix 2, Meristogenys amoropalamus (larval morphotype 3 -a): Shimada et al., 2007 b, p. 59–63. Diagnosis. This species is a member of the genus Meristogenys, based on the morphology of the larval specimens molecularly assigned to it (larval morphotype 3 -a in Shimada et al., 2007 a); these had diagnostic characteristics of the genus: abdominal sucker; divided upper jaw sheaths; and ribs on the outer surface of both jaw sheaths. Meristogenys dyscritus is a small species of the M. jerboa species group (Matsui, 1986), with male SVL 31.7–39.5 mm, female SVL 61.0– 71.2 mm; rear of thigh dark brown, dusted with small irregular light spots; web poorly developed leaving terminal two phalanges on fourth toe free of broad webbing; ova relatively small and heavily pigmented in the animal pole; upper third of iris sandy brown to yellowish brown. Etymology. The specific name is derived from dyskritos (Gr.), doubtful or indistinguishable, referring that the new species is difficult to distinguish from M. amoropalamus in adult morphology. Holotype. Osaka Museum of Natural History (OMNH) Am 8069, an adult male from Liwagu River, 1500 m, Headquarters of Kinabalu National Park, Sabah, Malaysia. Collected in March, 1979, by Masafumi Matsui. Paratypes. Two males and a female from the type locality; Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn (ZFMK) 41644; Sabah Parks (SP) 26481, 21392. Referred specimens. Forty-five males and 16 females from the type locality (Liwagu), Mahua, and Wario (See Appendix). Description of Holotype (measurements in mm). Body moderately slender, SVL 31.7; head subtriangular, slightly longer (12.8) than wide (12.7); snout blunt, projecting slightly beyond lower jaw; eyes elevated; canthi sharp, slightly concave; lores slightly oblique, concave; nostrils lateral, just below canthal edge, distinctly closer to tip of snout (2.4) than to eye (2.7); IND (4.3) wider than IOD (3.0); latter narrower than UEW (3.4); SL 5.2; pineal spot visible, slightly behind the line connecting anterior corners of orbits; tympanum distinct, TDv (3.5) and TDh (3.4), two-thirds of EL (5.2); T-EL (0.8) one-fourth of TDv and TDh; vomerine teeth obvious, in small oblique groups separated by the width of one group, groups on line connecting rear rims of choanae; tongue deeply notched, without papilla; paired subgular vocal sacs form gular pouches at corners of throat; vocal opening just inside commissures of jaws. Fingers slender, first and second subequal, much shorter than third; tips expanded into disks having circummarginal grooves; disks of all fingers subequal in diameter, one-third diameter of tympanum; no fringes of skin along fingers; no supernumerary metacarpal tubercles; distinct nuptial pads covering dorsal and medial surfaces of the first finger from its base to subarticular tubercle. Forelimb length unavailable; LAL 19.1; HAL 11.0; HLL 69.7; tibia long (23.1); heels overlapping when limbs are held at right angles to body; THIGH (19.9) and FL (20.3) much shorter than TL. Toe disks similar to those of fingers in shape and size; webbing not fully extending to disk on fourth toe, leaving two phalanges free of broad sheet; excision of web between fourth and fifth toes proximal to middle subarticular tubercle of fourth toe; a narrow fringe of skin along medial edge of first toe; inner metatarsal tubercle elliptical, shorter than distance between it and subarticular tubercle of first toe; a small round, raised outer metatarsal tubercle. Skin of dorsum finely granular on head and trunk; a weak fold from above eye to axilla; a low dorsolateral glandular fold; side of trunk coarsely granular; limbs strongly rugose above; throat smooth; chest and abdomen weakly rugose. Colour in life (See coloured pictures of Fig. 7). Dorsum light brown with small dark spots; lores with dark streak below canthus; upper lip light grey; a blackish brown band beginning behind eye bordering rear of the tympanum, diverging above the tympanum and nearly reaching the inguinal area, dorsal and ventral boundaries sharp; limbs marked dorsally with alternating light and dark brown crossbars, the darker ones narrower; a short dark streak ventrally at insertion of arm; rear of thigh light brown with scattered irregular light spots; throat and chest whitish with dots of melanophores; lower lip vaguely barred with blackish brown; ventral surface of tibia whitish, with heavy pigmentation of melanophores. Iris colour. Although iris colour of the holotype is unavailable, other specimens from the type locality (Liwagu) had the bicoloured iris, sandy brown above and reddish brown below. Colour in alcohol. Color pattern has not been changed even after preservation in ethanol for several years, except for iris colour which disappeared soon after the fixation in formalin solution. Eggs: Ova bicoloured, heavy pigmentations on the animal pole, cream white on the vegetal pole; clutch size 750–1500; ovum diameter 1.6–1.8 mm. Larvae (See coloured pictures of Fig. 8): We examined 67 specimens of stage 26–41 of Gosner (1960) from Mahua. These specimens are identical to those used by Shimada et al. (2007 a) to define their larval morphotype 3 - a. Head-body length (HBL) 10.0– 16.8 mm. Head-body oval, broadly rounded at snout, flat below, head-body width (HBW) maximum at level of spiracle 61–71 % (median= 65.8 %) of HBL; depth 49–64 % (median= 55.4 %) of HBW; eyes dorsolateral, not visible from below, pointing outward, eyeball 13–15 % (median= 13.7 %) of HBL; interorbital 123–169 % (median= 147.6 %) of eye diameter; eye-snout distance 36–46 % (median= 41.1 %) of HBL; nostril open, rim not raised, closer to eye than to tip of snout; internarial 57–80 % (median= 69.8 %) of interorbital. Oral disk ventral, width 57–73 % (median= 65.7 %) of HBW; upper lip separated from snout by a groove; upper lip with short marginal papillae in lateral third, inframarginal papillae near corner; lower lip with uninterrupted row of short marginal papillae; labial tooth row formulae 6 (4–6)/ 6 (1) in three specimens, 6 (4–6)/ 7 (1) in 48 specimens, and 6 (4–6)/ 8 (1) in 13 specimens; jaw sheaths heavy completely black except for outer margins covered by thin film; upper sheath film thicker than the lower; outer surface of lower jaw sheaths with several weak ribs; margin finely serrate, 3–11 and 4–7 serrae on a half of upper and lower jaw sheaths, respectively; upper jaw sheaths Mshaped, lower V-shaped; both jaw sheaths divided; a large suctorial abdominal disk following oral disk; peripheral part of disk darkened and keratinized; length 40–52 % (median= 46.5 %) of HBL; width 78–98 % (median= 87.9 %) of HBW. Spiracle sinistral; tube moderately long, length subequal to length of eyeball, pointing upward and backward, free of body wall for half its length; anal tube median, free of tail; tail heavily muscled, dorsal margin strongly convex, deepest before middle, tapering to slightly pointed tip; tail length 143–179 % (median= 164.7 %) of HBL, maximum depth 23–33 % (median= 26.5 %) of length; caudal muscle deeper than fins in basal half; dorsal fin origin behind body, fin deeper than ventral fin except in final fourth; ventral fin origin at end of proximal third of tail; head-body with four pairs of glandular clusters; a postorbital cluster about an eye length behind eye, with 0–3 glands; a infraorbital at the base of snout, with 1–3 glands; a prespiracular cluster just anterior to spiracle, with 0–4 glands; a midlateral at the posterior end of body, with 0–8 glands; no ventral and dorsal fin glands; 0–16 ventral fin glands; head-body scattered dorsally with minute protuberances anterior to eye in developed larvae; the area occupied by spinules and their density increasing with stage of development; lateral line pores indistinct. Head-body light brown dorsally and laterally, sometimes posterior half of lateral surface dark brown; caudal muscle light brown; translucent fins with scattered pigmentations. Other ten larval specimens from Bundu Tuhan (st. 28), Liwagu (st. 32), Poring (st. 31), and Wario (st. 30 and 32) had characteristics similar to Mahua specimens, but all ten specimens had six rows of lower labial teeth. Variation. Males from Mahua tended to be smaller than males from Liwagu and Wario in SVL (Table 3). Although relatively weak web development is one of the diagnostic characters of this species, some females from Liwagu and Wario had developed broad webs, which reached to the toe disk. Individuals usually had a bicoloured iris, sandy brown above and reddish brown below, but some females from Liwagu had monocoloured reddish brown iris. Larvae from Mt. Kinabalu tended to have fewer labial tooth rows in lower jaw than the specimens from Mahua (Crocker Range). In Mahua, only 4.7 % of larvae had six rows of labial teeth on the lower jaw, and all other larvae had seven or eight rows, whereas all ten larvae collected around Mt. Kinabalu (Bundu Tuhan, Liwagu, Poring and Wario) had six rows. Calls. We recorded calls of a male of M. dyscritus (SP 26472) at Liwagu at the midnight of 14 June 2007 (Fig. 9) at an air temperature of 17.5 C. The male perched on a low shrub, 50 cm above ground, and 1 m from the river. Many other males were also calling around there. The call was emitted sporadically with several minutes intervals, and consisted of a short, unpulsed note, lasting about 0.11 sec. A call included three continuous phases, of which the first one had a marked frequency modulation; it descended quickly in frequency from 8000–10000 Hz down to about 6000 Hz. In the second phase, the frequency was kept stable around 6000 Hz. The third phase had a remarkable frequency modulation again; it ascended quickly in frequency from 6000 up to 9000 Hz. Dominant frequencies were traced at about 6000–10000 Hz ranges, and harmonic bands were often found between 12000–15000 Hz ranges. Comparisons. Among 11 known species of Meristogenys, M. kinabaluensis (Inger, 1966), M. poecilus (Inger and Gritis, 1983), M. stenocephalus Shimada et al. 2011, M. stigmachilus Shimada et al. 2011, and M. whiteheadi (Boulenger, 1887) have a large body size (male SVL> 45 mm, female SVL> 70 mm) and are easily distinguished from M. dyscritus. From some small species, M. dyscritus is differentiated in the poorly developed toe webs [In M. jerboa (Gunther, 1872), M. macrophthalmus (Matsui, 1986), M. maryatiae Matsui et al., 2008, M. orphnocnemis (Matsui, 1986), and M. phaeomerus (Inger and Gritis, 1983), webs reach at the toe disk], heavy pigmentations on the ventral surface of tibia (M. jerboa, M. maryatiae, M. orphnocnemis, and M. phaeomerus have only scattered pigmentation), small irregular light spots on the rear of thigh (M. jerboa and M. macrophthalmus have large bright blotches), and black streaks around eyes, lores and tympanum (M. jerboa, M. maryatiae, M. orphnocnemis, and M. phaeomerus usually lack such colour patterns). Meristogenys dyscritus resembles M. amoropalamus in adult morphology, but the typical individuals can be identified through morphological characters. The upper iris of M. dyscritus is usually sandy brown, but is yellowish green in M. amoropalamus (Fig. 6), although this character changes after anaesthesia, and there might be an intraspecific variation. Additionally, females of M. dyscritus have smaller (Fig. 5) ova and larger clutch size (Fig. 4) than M. amoropalamus (See “Reproductive characters” in Result). In females, M. dyscritus have longer hand (usually longer than 31 % of SVL) than M. amoropalamus (usually less than 31 % of SVL) (Fig. 10). Males of M. amoropalamus usually have larger tympanum (more than 13 mm 2) than those of M. dyscritus (less than 15 mm 2), although there is a range of overlap (Fig 11). Contrary to the adult morphology, the larvae of M. dyscritus is quite different from those of M. amoropalamus in morphological characters. The larvae of M. dyscritus have six rows on upper jaws, while those of M. amoropalamus have seven. In the young developmental stages, the lower jaw sheaths of both larvae is divided, but only those of M. amoropalamus fuse into a large single jaw sheath before metamorphosis. Glands in a tail are limited to ventral fin in larval M. dyscritus, while they are present in both fins of larval M. amoropalamus. For the comparison between larval M. dyscritus and other Meristogenys larvae in Sabah, see the KEY TO MERISTOGENYS LAR- VAE FROM SABAH. The larvae of M. dyscritus share divided lower jaw sheaths, six rows of labial teeth on the upper jaw, and no glands on their dorsal fin or ventral surface with larval M. orphnocnemis. The larvae of M. dyscritus, however, have rectangular body shape (body oval in M. orphnocnemis), surface projections restricted to head (projections on head and body in advanced stage in M. orphnocnemis), and usually seven or eight rows of lower labial teeth (usually six rows in M. orphnocnemis; See Shimada et al., 2007 a for details). Range. Besides the type locality Liwagu, 1500 m a.s.l., adults of this species have been collected from Wario (900 m a.s.l.) in the Kinabalu National Park, and Mahua (1063 m a.s.l.) in the Crocker Range National Park, all in Sabah, Malaysia. The larvae of this species have also been collected from Bundu Tuhan (990 m a.s.l.) and Poring (500 m a.s.l.) in/around the Kinabalu National Park. Natural history. Gravid females were collected in July (Liwagu), August (Mahua and Wario), November (Wario), and December (Mahua). Larvae collected in March (Liwagu and Mahua), August (Liwagu, Wario, Bundu Tuhan, and Mahua), and November (Poring) showed a wide range of developmental stages. Thus, there seems to be no particular reproductive seasons. Larvae were collected from rivers with a width of 5– 10 m. At night, they make a mixed cluster with larvae of M. kinabaluensis, M. orphnocnemis, and Huia cavitympanum on top of flat rocks in the river.Published as part of Shimada, Tomohiko, Matsui, Masafumi, Yambun, Paul & Sudin, Ahmad, 2011, A survey of morphological variation in adult Meristogenys amoropalamus (Amphibia, Anura, Ranidae), with a description of a new cryptic species, pp. 33-56 in Zootaxa 2905 on pages 42-50, DOI: 10.5281/zenodo.20559
Leptobrachium waysepuntiense Hamidy & Matsui, 2010, sp. nov.
Leptobrachium waysepuntiense sp. nov. Holotype: MZB Amp 15862, an adult female collected among the leaf litter in primary forest in Way Sepunti trail (05°03’ 51 ’’S, 104 °02’05’’E, 691 m a.s.l.; Figure 1), Kubu Perahu village, District of Liwa, Lampung Province, Sumatra, Indonesia, collected by A. Hamidy, S. Kirono, D. Susanto, Marji, Andi, and Habsi at 20:00 h on 14 February 2009 (Figures 2, 3). Paratypes: MZB Amp 14592, an adult male collected by A. Ul Hasanah and W. Endarwin from near the type locality (05°03’ 51 ’’S, 104 °02’07’’E, 852 m a.s.l.) in November 2004; KUHE 42805, a juvenile, collected on 13 February 2009, collectors and locality same as for holotype. Etymology: The specific name refers to the locality where this species was collected. Diagnosis: The new species is placed in Leptobrachium by having the combination of: femoral glands present; oval, flat axillary glands present; inner palmar tubercle circular, not extending along first metacarpal; vomerine teeth absent; snout and/or dermal palpebral projections absent; rictal glands and ventrolateral glandular ridges absent; spines on the upper lip absent. A medium-sized Leptobrachium (adult female 58.3 mm and male 50.0 mm in SVL; Table 1); iris in adult light blue with fine black reticulations, but is light grey in juvenile; dorsum uniform dark grey; dark grey laterally with white and orange dots; top of head from interorbital to parietal region, and dorsal side of fingers and toes faintly orange; greyish on belly and brownish on throat with white dots particularly on chest; no markings at groin; femoral glands very small. Species Males Females Description of Holotype (measurements in mm): Body tapering to the groin (SVL 58.3), head broad and depressed, slightly longer (HL 25.5: 43.7 % of SVL) than wide (HW 24.4: 41.9 % SVL); snout rounded from above, truncate in profile, very slightly projecting beyond lower jaw; eye large and obviously projecting from sides of head, smaller (EL 8.6: 14.8 % SVL) than snout (SL 10.5: 18.0% SVL); canthi sharp, lores oblique, moderately concave; nostrils lateral, below canthus, distinctly closer to tip of snout (S-NL 4.8: 8.2 % SVL) than to eye (N-EL 6.0: 10.3 % SVL); internarial distance (IND 4.2: 7.2 % SVL) much shorter than interorbital distance, (IOD 8.1: 13.9 % SVL), latter wider than upper eyelid (UEW 7.5: 12.9 % SVL); no pineal spot; tympanum distinct, diameter (TD 5.0: 8.6 % SVL) about three-fifth that of eye and separated from eye by half of its diameter (T-EL 2.5: 4.3 %SVL); vomerine teeth absent; tongue heart-shaped, without papillae, notched posteriorly. Forelimb slender and long (FLL 43.2: 74.1 % SVL), about three-fifths of hindlimb; fingers moderately slender, unwebbed; first finger (FFL 6.6: 11.3 % SVL) slightly longer than fourth and second, third much longer (TFL 10.3: 17.7 % SVL); tips rounded, not swollen; inner palmar tubercle large (IPTL 2.8: 4.8 % SVL), not extending onto first metacarpal and smaller outer palmar tubercle (OPTL 2.5: 4.3 % SVL); subarticular tubercles indistinct, replaced by low callous tissue. Hindlimb slender and relatively short (HLL 71.0: 121.8 % SVL); heels not meeting when legs are held at right angles to body; tibia slightly longer (TL 20.3: 34.8 % SVL) than foot (FL 19.8: 34.0% SVL); tibiotarsal articulation of adpressed limb reaching to the middle of tympanum (Table 3); third toe longer than fifth; toe tips similar to those of fingers; toe webs poorly developed, webbing formula I 2 -- 2 + II 1 + -- 3 III 2 + -- 4 - IV 4 - -- 2 V (Figure 3 C); inner metatarsal tubercle low, oval, length (IMTL 2.6: 4.5 % SVL) more than half distance between tip of first toe and tubercle (1 TOEL 4.1:7.0%); outer metatarsal tubercle absent; subarticular tubercles obscure, but elongate, replaced by low callous tissue. Skin above nearly smooth, with minute granules scattered posteriorly, especially around waist; granules denser laterally on body and on posterior thigh; ventrally slightly granular; a very low supratympanic ridge from eye to behind tympanum; indistinct low of dermal ridges on upper surface of forelimb; a flat pectoral gland at median border of axilla behind arm insertion; minute femoral gland on posterior surface of thigh. Colour: In life, dorsally dark brownish grey with a faint V-shaped brownish orange interorbital and parietal marking (Figure 3 A); grey colour fading laterally to light grey on ventral side (Figure 3 B); laterally and ventrally dotted with white and orange, especially densely on sides; white dots denser on chest and throat than on abdomen; toes orange brown dorsolaterally; iris light blue with black reticulations; light blue orbital arc surrounding iris visible when eye opened maximally; no black bars around lips; supratympanic ridge bordered by a very thin brownish orange line; forelimb dorsally vaguely barred with dark brown; posterior thigh spotted with white and orange; no dark markings around groin from posterior flank to anterior thigh. In preservative, the aspects of the colour pattern remain, but the dorsal ground colour has been darkened and orange dots have faded to white. Variation: The male paratype (MZB Amp 14592) is morphologically similar to holotype, but has smaller body, fewer dark reticulations on blue iris, more rugose dorsal skin, more distinct dark crossbars on the dorsal sides of limbs, and more distinct markings between interorbital and on upper half of tympanum. It has an internal vocal sac and a pair of vocal sac openings. The juvenile paratype (KUHE 42805; Figure 4) has a light grey iris with irregular black reticulations, and a distinct orange line from canthus through margin of upper eyelid and above tympanum to arm insertion. Narrow dark brown bars each with orange spots medially are distinct on dorsal half of limbs, and toes are dorsally light brown. Comparisons: Small number of samples limited statistical comparisons of morphometric characters, but some dimensions did not overlap between L. waysepuntiense and other species (Table 2). The new species tended to have longer LAL and FL than L. hasseltii, longer FL than L. hendricksoni, and longer LAL, FL, and IMTL than L. abbotti, all relative to SVL. In addition, the new species seemed to have larger FL/TL ratio than L. nigrops (Table 2). In the new species, the tibiotarsal articulation reached the centre of the tympanum, but it barely reached the posterior end of the tympanum in many specimens of L. gunungense, and exceeded the anterior end of the tympanum in more than half specimens of L. montanum (Table 3). More discrete differences were found in qualitative characters (Table 4). Leptobrachium waysepuntiense differs in eye colour from all the other congeneric species including the subgenus Vibrissaphora. In most species of Leptobrachium from China and Indochina like L. chapaense, L. hainanense and L. huansen (Liu et al. 1973; Yang 1991; Dubois & Ohler 1998; Lathrop et al. 1998; Fei 1999; Fei et al. 2009), L. xanthospilum Lathrop, Murphy, Orlov & Ho and L. banae Lathrop, Orlov, Murphy & Ho, as well as species of Vibrissaphora, light blue or white colour is limited to the dorsal half of the iris, unlike L. waysepuntiense with totally light blue iris. Of the Indochinese species, L. buchardi has upper part of iris pale green (Ohler et al. 2004), and L. mauhoti has black iris surrounded by an orange-red crescent dorsally (Stuart et al. 2006). Leptobrachium hendricksoni from southernmost Thailand through Peninsular Malaysia, Sumatra to Borneo (Berry 1975; our own observations), L. pullum from southern Vietnam (Smith 1921) and L. smithi from Thailand to southern Malay Peninsula (Matsui et al. 1999) share dorsal half of the iris scarlet or yellow in colour. Leptobrachium nigrops from Peninsular Malaysia, Sumatra and Borneo (Berry 1975; our own observations) and all Bornean species, L. montanum, L. abbotti (Inger et al. 1995; Malkmus et al. 2002; our own observation), and L. gunungense (Malkmus et al. 2002; our own observations) have a totally black iris. The iris of L. hasseltii from Sumatra to Bali, once reported as scarlet (Iskandar 1998), is actually totally black (our observations). In addition to the iris colour, L. waysepuntiense differs from five of the seven congeners from Sundaland as follows; Leptobrachium waysepuntiense, without distinct dorsal markings, is differentiated from L. hasseltii and L. nigrops with distinct blotches on back, and L. smithi with confluent dark markings (Matsui et al. 1999). Furthermore, L. waysepuntiense clearly differs in ventral marking from L. hendricksoni with many black spots and L. abbotti with large dark lead-grey to black spots. The remaining two species, L. montanum and L. gunungense, with totally dark iris can be easily differentiated from L. waysepuntiense with light blue iris in life, but are otherwise very similar, although femoral glands are larger and more distinct in L. montanum than L. waysepuntiense (Figure 4 B). Species N Sex Beyond anterior edge Between anterior and Behind posterior edge of tympanum posterior edge of tympanum of tympanum Species Iris colour Marking at groin Femoral gland L. waysepuntiense all parts light blue with fine black reticulations no very small white spot L. hasseltii all parts black yes very large white blotch L. hendricksoni upper part orange yes large white blotch L. nigrops all parts black yes large white blotch L. montanum all parts black no small white spot L. abbotti all parts black no large white blotch L. gunungense all parts black no very small white spot L. smithi upper part yellow, orange, or scarlet yes medium-sized white spot Range: Southwestern Sumatra, Indonesia. Known from the type locality at Way Sepunti trail, Kubu Perahu, Liwa, West Lampung, but probably also occurs in western of Jambi Province, central Sumatra (Matsui et al., unpublished data, see below) (Fig. 1). Natural history: Larval, acoustic and other ecological data are unknown. Way Sepunti trail, where the type series of L. waysepuntiense was collected, is in primary forest and located approximately 250 m from a rocky stream. On this trail, the new species was found at a higher elevation (691–852 m a.s.l.) than where L. hasseltii occurred (300 m a.s.l.). Other species found on this trail are: Leptophryne borbonica Tschudi, Megophrys nasuta (Schlegel), Kalophrynus pleurostigma Tschudi, Limnonectes kuhlii (Tschudi), Odorrana hosii (Boulenger), Hylarana crassiovis (Boulenger), Huia sumatrana Yan g, Microhyla palmipes Boulenger, M. berdmorei (Blyth), and Rhacophorus sp. Neither eggs nor larvae were found and calls were not heard in mid February.Published as part of Hamidy, Amir & Matsui, Masafumi, 2010, A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia, pp. 34-44 in Zootaxa 2395 on pages 36-40, DOI: 10.5281/zenodo.19395
Maulisia argipalla Matsui & Rosenblatt 1979
Maulisia argipalla Matsui & Rosenblatt, 1979. Pitted Tubeshoulder. To 19.5 cm (7.7 in) SL (Sazonov et al. 1993). Atlantic and Pacific Oceans (Fricke et al. 2020); northern Pacific Japan (Balanov et al. 2009); Sea of Okhotsk; western Bering Sea near Karaginskiy Trench, eastern Bering Sea north of Semisopochnoi Island, and Gulf of Alaska (59°07’N, 146°54’W) (Stevenson et al. 2009) to Chile, including Gulf of California (Matsui and Rosenblatt 1987). Mesopelagic and bathypelagic; depth: 200–2,607 m (656–8,551 ft) (Porteiro et al. 2017).Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 50, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/557800
Rastrelliger faughni Matsui 1967
<p> 304. <i>Rastrelliger faughni</i> Matsui, 1967:74, figs.1, 5</p> <p>Paratype: USNM 76606 (2, 192–201), Takao, Taiwan, [South China Sea, 3–4 Dec. 1914, coll. F. Baker]; USNM 195321 (1, 202), [Albatross Expedition], Soo Wan Bay, Formosa, [South China Sea, 3–9 m, 29 Jan. 1910].</p> <p>Other type: USNM 190018 (holotype) and 28 paratypes.</p>Published as part of <i>Ho, Hsuan-Ching & Shao, Kwang-Tsao, 2011, 2957, pp. 1-74 in Zootaxa 2957</i> on page 6
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