A new from Borneo

SALAMANDRA 44 4 193-205 Rheinbach, 20 November 2008 ISSN 0036-3375

A new treefrog (Anura: : Rhacophorus) from Gunung Mulu, Borneo

J. Maximilian Dehling

Abstract. A new treefrog of the Rhacophorus is described from a small montane stream on the southern slope of Gunung Mulu in eastern Sarawak, Malaysian Borneo. The new species is distinguish- able from its Southeast Asian congeners by the combination of the following characters: small size; snout sharply pointed, forming a low swollen rostral tubercle at its anteriormost tip; supratympanic fold thick and conspicuous; tympanum diameter one-third of that of eye; canthus rostralis sharp; interorbital dis- tance greater than upper eyelid width and internarial distance; vomerine teeth in two oblique series; der- mal appendages on trunk and limbs absent; hands and feet not fully webbed; dorsum uniformly yellow- ish-green in life; iris ruby-coloured in life with a light grey margin; advertisement call consisting of three or four notes, with energy maximum at 4050-4380 Hz. Key words. Amphibia, Anura, Rhacophoridae, Rhacophorus penanorum sp. n., , natural his- tory, advertisement call, Gunung Mulu National Park, Sarawak, Malaysia, Borneo.

Introduction During field work in the park I collected specimens of a species of Rhacophorus Kuhl Gunung Mulu National Park (3°56’-4°6’N, & van Hasselt, 822 that does not match any 4°47’-5°00’E) is located in eastern Sara- of the described species of the genus from wak on Borneo, the largest of the Greater Borneo and from any other part of Southeast Sunda Islands. It is named after Gunung (= Asia. I therefore describe it herein as new to “Mount”) Mulu, the second highest peak in science. The treefrog genus Rhacophorus is Sarawak (2376 m elevation), and encompass- distributed from India to Japan and south- es an area of 544 km2 (Hazebroek & Kash- wards to the Indo-Malayan archipelago and im bin Abang Morshidi 2006). Situated the Philippines and currently contains 76 within the borders of the national park are species (Frost 2007, Bordoloi et al. 2007, the type localities of fourteen species, Dehling & Grafe 2008). Sixteen species eight of which have not been recorded else- have been reported from Borneo (Inger & where (Dring 983a, 983b, 987, Kiew 984, Tan 996, Das & Haas 2005, Dehling & Dubois 987, Inger et al. 995). Neverthe- Grafe 2008). less the herpetofauna of the park has hardly been explored. Most of the field work con- ducted so far took place during the joint ex- Materials and methods pedition of the Sarawak Government and the Royal Geographic Society between 977 and Type specimens were collected on 4 January 978, in the course of which 7 species of an- 2008 between 945 and 2000 h, kept alive for urans were recorded (Dring 978, Dring & two days, euthanised using a chlorobutanol Kiew 982). In total 76 species of amphibi- solution, fixed and preserved in 70% ethanol, ans (Dring & Kiew 982, Inger & Stuebing and measured four weeks after collection. 992, Sanchez-Herraiz et al. 995, Malk- The following measurements were taken mus 2002) and 9 species of reptiles (Das et with a digital caliper (to the nearest 0. mm): al. 2008) have been reported from Gunung snout-vent length (SVL, from tip of snout Mulu National Park so far. to vent); tibia-fibula length (TFL, measured

© 2008 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT) http://www.salamandra-journal.com 193 J. Maximilian Dehling with both knee and tibio-tarsal articulation ger & Stuebing (2005), Wilkinson et al. flexed); head width (HW, distance between (2005), Das & Haas (2005), Matsui & Pan- angles of jaw); head length (HL, distance ha (2006), Ohler & Delorme (2006), and from angle of jaw to tip of snout); horizon- Bordoloi et al. (2007). tal eye diameter (ED); horizontal tympanum Advertisement calls from the type speci- diameter (TD); upper eyelid width (EW); in- mens were recorded in the habitat using a terorbital distance (IO, shortest distance be- Sony WM-D6C stereo cassette recorder and tween upper eyelids); eye to nostril distance a Sony ECM-S959C microphone. Call re- (EN, distance between anterior margin of eye cordings were digitised at 6 bits and 44 kHz and nostril); nostril to snout distance (NS, and analysed using Syrinx 2.6h sound analy- distance between nostril and tip of snout); sis program (John Burt, www.syrinxpc.com) internarial distance (NN, distance between and Adobe Audition .5 software. nostrils); hand length (HND, distance from base of thenar tubercle to tip of third finger); foot length (FOT, distance from base of in- Systematics ner metatarsal tubercle to tip of fourth toe). Rhacophorus penanorum sp. n. The webbing formulae are given as proposed (Figs. -7) by Myers & Duellman (982). Notes on the colouration in life were taken from digital Holotype: ZMB 7078, adult male, from a photographs taken in the habitat and during small stream, 45 minutes by foot below camp two days after collection. four on the southern flank of Gunung Mulu, For comparisons I examined museum approx. 650 m elev., Gunung Mulu National specimens of several species of Rhacopho- Park, Miri Division, Sarawak, Malaysia, col- rus (see Appendix). Museum abbreviations lected by J. M. Dehling, 4 January 2008. are as follows: Field Museum of Natural His- tory, Chicago, Illinois, USA (FMNH), Mu- Paratopotypes: ZMB 7079, 70720, two adult seum und Forschungsinstitut Senckenberg, males, collected with the holotype. Frankfurt am Main, Germany (SMF), Sabah Parks Zoological Museum, Kinabalu Park Diagnosis: A species of the genus Rhacoph- Headquarters, Ranau District, Sabah, Malay- orus, distinguishable from its congeners by sia (SP), Zoological Museum of the Depart- the combination of the following characters: ment of Biology, Universiti Brunei Darussal- am, Bandar Seri Begawan, Brunei Darussal- am (UBD), Zoologisches Museum der Hum- boldt-Universität zu Berlin, Berlin, Germany (ZMB). Additional information about char- acters used to compare Southeast Asian spe- cies of Rhacophorus were taken from: Bou- lenger (882), Ahl (927), Smith (930), Wolf (936), Inger (954), Taylor (962), Inger (966), Brown & Alcala (994), Manthey & Grossmann (997), Manthey & Steiof (998), Inger et al. (999), Ohler et al. (2000), Ziegler & Köhler (200), Or- lov et al. (200), Chanda (2002), Ziegler (2002), Harvey et al. (2002), Malkmus et Fig. 1. Holotype of Rhacophorus penanorum sp. al. (2002), Wilkinson & Rao (2004), Mat- n. (ZMB 70718) in life. Photograph taken in the sui (2005), Inger & Iskandar (2005), In- habitat.

194 A new Rhacophorus from Borneo

() small size (SVL of adult males 33.6-34.2 mm); (2) snout sharply pointed, projecting beyond mandible, sloping forward from nos- trils then backward to mouth, forming a low swollen rostral tubercle at its anteriormost tip; (3) head wider than long and wider than body; (4) supratympanic fold thick and con- spicuous; (5) tympanum diameter one-third of that of eye; (6) canthus rostralis sharp; (7) interorbital distance greater than upper eye- lid width and internarial distance; (8) vomer- ine teeth in two oblique series; (9) dermal appendages on trunk and limbs absent; (0) Fig. 3. Lateral view of the head of a paratype hands and feet not fully webbed; () dor- of Rhacophorus penanorum sp. n. (ZMB 70719). sum uniformly yellowish-green in life; (2) Note the brownish-green daytime-colouration of iris ruby-coloured in life with a light grey the dorsum. margin; (3) advertisement call consisting of three or four notes, with energy maximum at 4090-4380 Hz.

Description of the holotype: Body moder- ately slender with a narrow waist; head short (HL/SVL 0.3), slightly wider than trunk and wider than long (HW/HL .23); snout project-

Fig. 4. Type series of Rhacophorus penanorum sp. n. (top to bottom: ZMB 70719, 70718, 70720), showing the variation in daytime-colouration.

Fig. 2. Ventral view of a paratype of Rhacophorus Fig. 5. Paratype of Rhacophorus penanorum sp. penanorum sp. n. (ZMB 70719) in life. n. (ZMB 70720) in life. Note the greyish-green daytime-colouration.

195 J. Maximilian Dehling ing beyond mandible, pointed in dorsal view, and IV; webbing formula of the hand I2.75- sloping forward from nostrils then back to- 2.75II.75-3III2-2-IV (Fig. 6); thenar tuber- ward mouth in profile, forming a low swollen cle oval, almost as large as base of Finger I rostral tubercle at its anteriormost tip (cf. Fig. (length 2.4 mm); inner and outer palmar tu- 3); canthus rostralis distinct, sharp, curved in bercles oval, low and indistinct, fused at their profile, slightly concave in dorsal view; loreal bases; narrow dermal ridge on postaxial edge region sloping, almost vertical; nostril about of Finger IV between base of disk and outer as close to eye as to tip of snout (EN/NS 0.99); palmar tubercle; metacarpals with several su- internarial distance greater than eye to nostril pernumerary small tubercles; nuptial pads or distance (NN/EN .43); eyes directed antero- asperities absent. laterally, protruding, large (ED/HL 0.45); pu- Legs slender, moderately long (TFL/SVL pil horizontal; eye diameter greater than eye- 0.53); heels slightly overlap each other when to-nostril distance (ED/EN .63); interorbital legs are folded right angle to body; tibio-tarsal distance wider than upper eyelid (IO/EW articulation reaching tip of snout; heel with- .30) and greater than internarial distance out dermal appendages; tarsus with several (IO/NN .08); tympanum visible, separated very small, indistinct white tubercles; relative from eye by distance subequal to its horizon- length of toes: I < II < III < V < IV; disks of tal diameter; tympanum diameter about one- toes smaller than those of fingers; subarticu- third of eye diameter (TD/ED 0.32); tym- lar tubercles numbering one on Toes I and II, panic annulus visible; skin not co-ossified to two on Toes III and V, and three on Toe IV; forehead; maxillary teeth present; choanae pedal webbing formula I.5-2II+-2.5III.5- located far laterally, at margins of roof of the 2.5IV2-+V (Fig. 7); inner metatarsal tubercle mouth; dentigerous processes of vomers low, oval, low and small (length .3 mm), no outer located near anterior edges of choanae, di- one; few supernumerary small tubercles on rected posteromedially, bearing small teeth, metatarsals; dermal ridge on postaxial edge separated from each other by distance equal of Toe V from disk to base of metatarsus. to four-thirds their length; tongue broad, bi- Measurements (in mm; holotype, followed furcated at base and free for about two-thirds by paratypes [ZMB 7079, 70720, respective- of its length; median lingual process absent; ly]): SVL 33.6 (34.0, 34.2), TFL 7.7 (7.6, 7.), median subgular vocal sac present. HW 2.6 (2.5, 2.5), HL 0.2 (0.2, 0.3), ED Dorsal surfaces, ventral side of limbs, 4.6 (4.9, 5.0), TD .5 (.6, .7), EW 3.3 (3.3, chin, throat, and chest region smooth; abdo- 3.4), IO 4.3 (4.4, 4.5), EN 2.8 (2.9, 2.8), NS 2.8 men coarsely granular; supratympanic fold (2.9, 3.0), NN 4.0 (4.0, 4.0), HND 0.8 (.4, thick and conspicuous, extending from pos- 0.9), FOT 4. (4.6, 4.). terior edge of upper eyelid to scapula region, overlapping tympanum for about one-fifth Colouration in life: Dorsum uniformly yel- on upper edge; supracloacal region unorna- lowish-green (Fig. ). Sides of head below mented, without ridge, flap or fold; several canthus rostralis and supratympanic fold, in- small tubercles in infraanal region and on cluding annulus of tympanum, slightly light- ventral side of thigh; small, indistinct white er green. Dorsal colouration fading to white tubercles along postaxial edge of forearm. on flanks. Chin, throat, and chest completely Arms moderately slender; tips of fingers white. Venter white, sparsely speckled with enlarged into broad oval disks, each with faint, reddish spots. Large black and dark circummarginal groove; disk of third fin- brown spots present on flanks. Groin region, ger wider than tympanum diameter; relative ventral sides of arm, thigh, tibia, hand and length of fingers: I < II < IV < III; subarticular foot, preaxial side of tarsus and dorsal side tubercles rounded, well developed, number- of Fingers I and II and Toes I-III largely un- ing one on Fingers I and II, two on Fingers III pigmented, appearing greyish violet (cf. Fig.

196 A new Rhacophorus from Borneo

2). Webbing between fingers translucent light tained its night colouration albeit somewhat brown with some greenish-white mottling. paler (Fig. 4). In the greyish-green paratype Webbing between toes darker with a con- even the iris turned completely grey during spicuous narrow green stripe between Toes the day (Fig. 5). In preservative, the paratypes IV and V. Iris ruby-coloured with a light grey resemble the holotype in colouration. margin. Advertisement call: In total, twelve advertise- Colouration in preservative: Dorsum colour- ment calls from the three type specimens were ation faded to greyish brown. Darker, narrow recorded prior to collection. The called crosslines faintly visible on dorsal surfaces of at irregular intervals between 30 secs and  thigh and tibia. Ventral side of arm, thigh, min. The call is a very short trill. Measure- and tibia changed to white. Small light brown ments of the recorded calls are given in Tab. blotches present on margin of lower jaw, . Air temperature was 6. °C. There was no chest, lateral sides of arm, and ventral surface significant frequency modulation within the of thigh. Colour of iris faded to bluish-grey. notes of a single call and there were no dif- ferences in energy maximum between three Variation: The paratypes are very similar to note and four note calls. Dominant frequen- the holotype in measurements and propor- cy spectrum of the notes lay between 3900 tions. They likewise have median subgular and 4600 Hz (Fig. 8). The intervals between vocal sacs and lack nuptial pads and asperi- the amplitude maxima of the notes ranged ties. There is little variation in the extent of between 34 and 43 ms (Fig. 9). webbing. In one paratype (ZMB 70720), the Apart from R. gadingensis, the call of webbing on the preaxial side of Finger IV which is unknown, all Bornean species of reaches only to the middle of the distal subar- Rhacophorus can be distinguished from R. ticular tubercle (2IV), in the other one (ZMB penanorum by their calls (Malkmus et al. 7079), the webbing on the preaxial sides of 2002, own data). Toes III and IV reaches to just below the dis- tal subarticular tubercles (2.25III.5-2.25IV). Distribution and ecological notes: The new Colouration in life was also similar to that of species is known only from the type locali- the holotype. During the night, both speci- ty, a small permanent headstream of Sungai mens were yellowish-green dorsally like the Tapin on the southern flank of Gunung Mulu holotype. However, one paratype turned (approx. 650 m elev.), 45 minutes by foot brownish-green during the day, the other one below camp four. The streambed consists of greyish-green, whereas the holotype main- rock in steep, fast-flowing sections and gravel

Tab. 1. Characteristics of the advertisement call of Rhacophorus penanorum sp. n. Measurements are given as mean ± standard deviation.

7078 7079 70720 Holotype Paratype Paratype number of three note calls recorded 2 5 – number of four note calls recorded  3  duration of three note call [ms] 98 ±  05 ± 3 – (range) (97-99) (0-0) duration of four note call [ms] 42 40 ± 0 3 (range) (29-48) energy maximum of notes [Hz] 440 ± 82 490 ± 88 4220 (range) (4050-4260) (4090-4380) (480-4270)

197 J. Maximilian Dehling

Fig. 6. Ventral view of the right hand of the ho- Fig. 7. Ventral view of the left foot of the holotype lotype of Rhacophorus penanorum sp. n. (ZMB of Rhacophorus penanorum sp. n. (ZMB 70718). 70718). in sections where the current is slower and new species is situated on one of their tradi- the stream forms small, shallow pools. The tional hunting trails. site is also described in Dring (983a, 983b). All specimens were found calling from leaves Comparisons: In the following Southeast overhanging the stream at heights between Asian congeners the webbing reaches the disk .5 and 2 m. The following species were found on the postaxial side of the third finger (f), on sympatric with the new species: Ansonia preaxial and postaxial sides of the fourth toe hanitschi Inger, 960, A. torrentis Dring, (t) or both (f & t), thus readily distinguishing 983, Leptobrachella brevicrus Dring, 983, these species from Rhacophorus penanorum: Xenophrys dringi (Inger, Stuebing & Tan, Rhacophorus annamensis Smith, 924 (f & t); 995), Limnonectes kuhlii (Tschudi, 838) - R. dennysi Blanford, 88 (t); R. dulitensis complex, Meristogenys kinabaluensis (Inger, Boulenger, 892 (t); R. exechopygus Inger, 966), Staurois natator (Günther, 858), Orlov & Darevsky, 999 (f & t); R. fasciatus and Philautus mjobergi Smith, 925. Dring Boulenger, 895 (f & t); R. feae Bouleng- (983b) reported Leptolalax sp. (= L. dringi er, 893 (f & t); R. georgii Roux, 904 (f & t); Dubois, 987) from the site. R. harrissoni Inger & Haile, 959 (f & t); R. htunwini Wilkinson, Thin, Lwin & Shein, Etymology: The species epithet is a patronym 2005 (f & t); R. jarujini Matsui & Panha, honouring the Penan people who live in the 2006 (t); R. kio Ohler & Delorme, 2006 (f Gunung Mulu area. The type locality of the & t); R. maximus Günther, 858 (f & t); R.

198 A new Rhacophorus from Borneo nigropalmatus Boulenger, 895 (f & t); R. pardalis Günther, 858 (f & t); R. poecilono- tus Boulenger, 920 (t); R. prominanus Smith, 924 (t); R. reinwardtii (Schlegel, 840) (f & t); R. rhodopus Liu & Hu, 960 (f & t); R. robinsonii Boulenger, 903 (f & t); R. rufipes Inger, 966 (f & t). Webbing between the fingers is confined to the bases in R. du- gritei (David, 872), R. everetti Boulenger, 894, R. hungfuensis Liu & Hu, 96, and R. kajau Dring, 983. The following Southeast Asian species of Rhacophorus differ from the new species in having dermal flaps or fringes Fig. 8. Sound spectrogram of a four-note call of a along the postaxial edge of the forearm (F), paratype of Rhacophorus penanorum sp. n. (ZMB the tarsus (T), or both (F & T): Rhacopho- 70720) given at 16.1°C. rus achantharrhena Harvey, Pemberton & Smith, 2002 (F & T); R. appendiculatus (Günther, 858) (F & T); R. baluensis In- ger, 954 (F & T); R. barisani Harvey, Pem- berton & Smith, 2002 (F & T); R. bipunc- tatus Ahl, 927 (F & T); R. calcaneus Smith, 924 (T); R. edentulus Müller, 894 (F & T); R. hoanglienensis Orlov, Lathrop, Murphy & Ho, 200 (F); R. margaritifer (Schlegel, 837) (F & T); R. monticola Boulenger, 896 (F & T); R. taronensis Smith, 940 (F & T); R. turpes Smith, 940 (F & T). The remain- ing Southeast Asian Rhacophorus species Fig. 9. Oscillogram of a three note call of the differ in the following characters from the holotype of Rhacophorus penanorum sp. n. (ZMB new species (characters given in parenthe- 70718) given at 16.1°C. ses). Rhacophorus belalongensis Dehling & Grafe, 2008, R. bimaculatus (Peters, 867), Swan & Grosjean, 2000 is considerably R. gadingensis Das & Haas, 2005, and R. larger, SVL of the male holotype being 6.5 gauni (Inger, 966) are somewhat smaller mm (SVL of males 33.6-34.2 mm), has a tym- species and have brown or grey dorsal col- panum diameter that is two-thirds of that of ouration (green), dorsal markings consisting eye (one-third of eye diameter), and a gold- of a brown interorbital band and crossbars en iris (red with a grey margin). Rhacophorus on thigh, tibia, tarsus and forearm (markings modestus Boulenger, 920, is brown dor- absent), a more or less rounded, obtuse snout sally (green), has a tympanum diameter wid- (pointed) and a large pointed tubercle at the er than half the eye diameter (one-third of tibio-tarsal articulation (absent). Rhacopho- eye diameter) and the fingers only one-third rus dorsoviridis Bourret, 937 has shorter webbed (webbing between fingers III and IV legs with TFL/SVL 0.40-0.42 (0.50-0.53) and to distal subarticular tubercles). Rhacophorus the tibio-tarsal articulation extending only to omeimontis Stejneger, 924 is larger, SVL of the posterior margin of the eye (extending to males up to 67.2 mm (up to 34.2 mm), has tip of snout), a larger tympanum with TD/ED a rounded snout (pointed), and a large tym- 0.52 (0.32-0.34), and a rounded snout (point- panum that is 70% of eye diameter (32-34%). ed). Rhacophorus duboisi Ohler, Marquis, Rhacophorus cyanopunctatus Manthey &

199 J. Maximilian Dehling

Steiof, 998 and R. orlovi Ziegler & Köh- ally with several large, darker brown blotches ler, 200 are greyish to reddish brown dor- (green without blotches), and a dark brown sally in life, with darker blotches on the dor- interorbital band (absent). Forearm, thigh, sum and crossbars on the limbs (dorsum uni- tibia and tarsus are dark-banded (without formly yellowish-green without markings), pattern), flanks, lateral surfaces of thigh, tib- have blue blotches on the flanks and later- ia and tarsus are yellow with dark blotches al surfaces of the thighs (flanks white with (white, dark-blotched flanks, lateral surfac- brown or black blotches, thighs unblotched), es of legs unpigmented), and the iris is red and have more extensively webbed toes, for- in the lower two-thirds and blue in its upper mulae being I-.5II-(.5-2)III-2IV.5-V third (red with a light grey margin). In most and I-.5II-.5III-2-IV.5-V, respective- specimens, one or several white infraocu- ly (versus I.5-2II+-[2.25-2.5]III.5-[2.25- lar spots are present (absent). Furthermore, 2.5]IV2-+V). In addition, R. orlovi has more the snout is pointed, but does not form a extensively webbed outer fingers, formula low swollen rostral tubercle at its anterior- being II+-2III.5-.5IV (versus II.75-3III2- most tip (forms a tubercle), and the eyelid di- 2(-)IV), and R. cyanopunctatus lacks vomer- ameter almost equals the interorbital width ine processes and teeth (present). with IO/EW .03-.2, mean .08 ± 0.04, n In males of Rhacophorus catamitus Har- = 5 (IO/EW .30-.33). Vomerine teeth are vey, Pemberton & Smith, 2002, the dor- present, but the dentigerous processes of the sum is brown with irregular darker brown vomers are shorter and separated from each blotches on the back and darker bands on the other by a distance equal to about twice their limbs (dorsum uniformly yellowish-green), length (longer, separated by four-thirds their the groin and ventral surfaces of the thighs, length). The advertisement call is similar in hands, and feet are orange and the iris is dirty frequency (maximum at 4050-4570 Hz), but yellow with a tinge of green in life (groin and typically consists of only two notes (Malk- ventral surfaces unpigmented, iris red with mus et al. 2002, own unpublished data). The a grey margin), webbing is less extensive average call duration is 92 ± 4 ms (n = 6 with hand webbing formula being II(2+-2.5)- calls from 3 individuals, air temperature 8.4 3.5III2.75-(2+-2.75)IV (versus II.75-3III2-2(-) °C; versus 03 ± 5/39 ± 8 ms for three note/ IV), and pedal webbing formula being I.75- four note calls, air temperature 6. °C); an (2+-2.5)II(.25-.75)-(2.75-3)III(.5-.75)-(2.5- exceptional call consisting of three notes had 3.25)IV(2.25-2.75)-(.25-.75)V (versus I.5- a duration of 253 ms (unpublished data). 2II+-[2.25-2.5]III.5-[2.25-2.5]IV2-+V), and a subtriangular tubercle is present at the ti- Remarks: The classification of Asian rhacoph- bio-tarsal articulation (absent). orids is still unstable, and species are trans- Rhacophorus penanorum is similar to R. ferred from one genus to another frequent- angulirostris Ahl, 927, a species that has ly (e.g. Wilkinson et al. 2002, Delorme been collected at Gunung Kinabalu, Gunung et al. 2005, Frost et al. 2006). The genus Trusmadi, and the Crocker Range in Sabah/ Rhacophorus might be paraphyletic with re- Borneo and at Padang on Sumatra (Inger spect to Tschudi, 838 or even 966, Inger & Tan 990, Inger et al. 2000, polyphyletic, and is in need of a comprehen- specimens listed in appendix). Both species sive taxonomic revision (Wilkinson et al. differ strongly in colouration (colouration in 2002, Frost et al. 2006). The new species is life of R. angulirostris is described in Mal- allocated to Rhacophorus because it shows kmus 992, 993, 995, Hoffmann 995, In- the following characters considered diagnos- ger et al. 996, Malkmus et al. 2002, Inger tic for the genus (Brown & Alcala 994, & Stuebing 2005): Males of R. angulirostris Manthey & Grossmann 997, Wilkinson & possess a tan to sandy brown dorsum, usu- Drewes 2000; see also Harvey et al. 2002):

200 A new Rhacophorus from Borneo

fingers webbed, skin of head free from skull, faces of thigh and groin golden yellow in life and outer two metacarpals separated by a (dorsal side of upper arm green, ventral side skin membrane. Disregarding these charac- of upper arm, lateral surfaces of thighs and ters, the new species might be confused with groin unpigmented in life), undersides of a Bornean member of the genus Philautus thigh yellowish (unpigmented). Gistel, 848, P. erythrophthalmus Stuebing & Wong, 2000. This species was described from 550 m elev. at Mt. Muruk Mio in Sabah Acknowledgements and resembles R. penanorum in having a light green dorsum, a “deep reddish orange” Thanks to Donny Andrew (K. Long Iman, Sara- iris, dark reticulate markings on the flanks, a wak) for guiding me to the type locality and for the tympanum diameter that is one-third of the companionship during the trip. Mohd. Shabu- din Sabki, Uning Paun (Sarawak Forest Depart- eye diameter, and in the absence of dermal ment), and Taha Wahab (Sarawak Forestry Cor- appendages on the limbs (Stuebing & Wong poration) kindly assisted in obtaining permits to 2000). Based on its small size, the absence of conduct research on in Gunung Mulu vomerine teeth, the absence of the musculus National Park (No. NPW.907.4.2(II)-76, dated cutaneus pectoris, and the m. geniohyoideus 03.09.2007) and to export collected specimens pars medials being free, the species was as- (No. 07437, dated 07.0.2008). Both permits were signed to the vermiculatus group of the ge- issued by Haji Ali bin Yusop. Gunther Köhler nus Philautus by Stuebing & Wong (2000). (SMF), Maklarin bin Lakim and Paul Imbun Taking into consideration that the most re- (SP), T. Ulmar Grafe (UBD), and Rainer Gün- ther and Detlef Langer (ZMB) provided wor- cent diagnosis of Philautus characterises the king space and let me examine the material under genus only by “aerial direct development of their care. Harold K. Voris and Alan Resetar eggs into froglets, without going through an (FMNH) sent comparative material of Rhacopho- aquatic tadpole stage” (Bossuyt & Dubois rus, which Mark-Oliver Rödel (ZMB) helped 200) and that the reproductive mode of P. to obtain. T. Ulmar Grafe (UBD) and Edgar erythrophthalmus, which is known only from Lehr (SNSD) read the manuscript. Bryan Stu- a single female, is unknown, the question is art (FMNH) drew my attention to the new spe- whether this species could possibly be con- cies’ similarity to Philautus erythrophthalmus. In- specific with R. penanorum, irrespective of draneil Das (UNIMAS) sent photographs of the type specimens of Rhacophorus gadingensis and the generic affiliation. However, it differs commented on the manuscript. Thanks also to from the new species (characters in paren- Brian Clark and the staff of the Gunung Mulu theses) in the following characters: small size, National Park for providing excellent logistic sup- the only specimen known, a mature female, port. Field work was supported by a grant from the having a SVL of only 26.2 mm (larger, SVL of German Academic Exchange Service (DAAD). adult males being 33.6-34.2 mm), snout blunt, abruptly truncate in profile (snout sharply pointed, projecting beyond mandible), nos- trils situated near the end of the snout, above References the symphysis (situated half-way between Ahl, E. (927): Zur Systematik der asiatischen eye and tip of snout), vomerine teeth absent Arten der Froschgattung Rhacophorus. – Sit- (present), Fingers III and IV webbed only at zungsberichte der Gesellschaft naturforschen- the base (webbed to distal subarticular tuber- der Freunde zu Berlin, 927: 35-47. cles), toes only half-webbed, formula being + + + + + Bordoloi, S., T. Bortamuli & A. Ohler (2007): I2 -2 II2 -3 III2-3IV3-2V (versus I.5-2II - Systematics of the genus Rhacophorus (Amphi- + [2.25-2.5]III.5-[2.25-2.5]IV2- V), Toe III bia, Anura): identity of red-webbed forms and longer than Toe V (Toe V longer than Toe description of a new species from Assam. – Zo- III), upper arm, anterior and posterior sur- otaxa, 653: -20.

201 J. Maximilian Dehling

Bossuyt, F. & A. Dubois (200): A review of the management and development plan. – Royal frog genus Philautus Gistel, 848 (Amphibia, Geographic Society, London. 345 pp. Anura, Ranidae, ). – Zeylanica, Dubois, A. (987): Miscellanea taxinomica batra- 6: -2. chologica (). – Alytes, 5: 7-95. Boulenger, G.A. (882): Catalogue of the Batra- Frost, D.R. (2007): species of the chia Salientia s. Ecaudata in the collection of world 5., an online reference. – American Mu- the British Museum. Second Edition. – Taylor seum of Natural History, New York, USA. Elec- and Francis, London. xvi+503 pp., 30 pls. tronic database accessible at http://research. Brown, W.C. & A.C. Alcala (994): Philippine amnh.org/herpetology/amphibia/index.php frogs of the family Rhacophoridae. – Procee- (.02.08). dings of the California Academy of Science, Frost, D.R., T. Grant, J. Faivovich, R.H. Bain, 48(0): 85-220. A. Haas, C.F.B. Haddad, R.O. de Sá, A. Chanda, S.K. (2002): Hand book of Indian am- Channing, M. Wilkinson, S.C. Donnellan, phibians. – Zoological Survey of India, Kolka- C.J. Raxworthy, J.A. Campbell, B.L. Blot- ta, India. 335 pp. to, P. Moler, R.C. Drewes, R.A. Nussbaum, J.D. Lynch, D.M. Green & W.C. Wheeler Das, I. & A. Haas (2005): A new species of Rhaco- (2006): The amphibian tree of life. – Bulletin phorus (Anura: Rhacophoridae) from Gunung of the American Museum of Natural History, Gading, Sarawak. – The Raffles Bulletin of Zo- 297: -370. ology, 53: 257-263. Harvey, M.B., A.J. Pemberton & E.N. Smith Das, I., B. Clark, S. Clark & E. McArthur (2002): New and poorly known parachuting (2008): An inventory of reptiles of Gunung frogs (Rhacophoridae: Rhacophorus) from Su- Mulu National Park, Sarawak, Malaysia (Bor- matra and Java. – Herpetological Monographs, neo). – Sarawak Museum Journal, 63(84): 27- 6: 46-92. 67. Hazebroek, H.P. & A. Kashim bin Abang Dehling, J.M. & T.U. Grafe (2008): A new tree- Morshidi (2006): National Parks of Sarawak. frog of the genus Rhacophorus (Anura: Rhaco- – Natural History Publications (Borneo), Kota phoridae) from Brunei Darussalam (Borneo). Kinabalu. xii+503 pp. – Salamandra, 44: 0-2. Hoffmann, P. (995): Untersuchungen zur An- Delorme, M., A. Dubois, S. Grosjean & A. Oh- urenfauna des Mt. Kinabalu (Borneo). Teil II: ler (2005): Une nouvelle classification gene- Familien Ranidae, Rhacophoridae. – Sauria, rique et subgenerique de la tribu des Philautini 7(3): 9-7. (Amphibia, Anura, Ranidae, Rhacophoridae). – Bulletin Mensuel de la Société Linnéenne de Inger, R.F. (954): Systematics and zoogeography Lyon, 74: 65-7. of Philippine Amphibia. – Fieldiana: Zoology, 33: 83-53. Dring, J. (978): Preliminary report. Gunong Inger, R.F. (966): The systematics and zoogeo- Mulu National Park frogs. – Unpublished re- graphy of the Amphibia of Borneo. – Fieldiana: port, 4 pp. Zoology, 52: -402. Dring, J. (983a): Frogs of the genus Leptobrachel- Inger, R.F. & D.T. Iskandar (2005): A collection la (Pelobatidae). – Amphibia-Reptilia, 4: 89- of amphibians from West Sumatra, with de- 02. scription of a new species of Megophrys (Am- Dring, J. (983b): Some new frogs from Sarawak. phibia: Anura). – The Raffles Bulletin of Zoo- – Amphibia-Reptilia, 4: 03-5. logy, 53: 33-42. Dring, J. (987): Bornean treefrogs of the genus Inger, R.F. & R.B. Stuebing (992): The monta- Philautus (Rhacophoridae). – Amphibia-Rep- ne amphibian fauna of northwestern Borneo. tilia, 8: 9-47. – Malayan Nature Journal, 46: 4-5. Dring, J.C. & B.H. Kiew (982): Frogs recorded Inger, R.F. & R.B. Stuebing (2005): A field guide within Gunung Mulu National Park. – p. 7 in to the frogs of Borneo (Second Edition). – Na- Anderson, J.A.R., A.C. Jermy & The Earl of tural History Publications (Borneo), Kota Ki- Cranbrook: Gunung Mulu National Park: a nabalu. vii + 204 pp.

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Inger, R.F. & F.L. Tan (990): Recently discovered Manthey, U. & C. Steiof (998): Rhacophorus and newly assigned frog larvae (Ranidae and cyanopunctatus sp. n. (Anura: Rhacophori- Rhacophoridae) from Borneo. – The Raffles dae), ein neuer Flugfrosch von der Malaii- Bulletin of Zoology, 38: 3-9. schen Halbinsel, Sumatra und Borneo. – Sau- Inger, R.F. & F.L. Tan (996): Checklist of the ria, 20(3): 37-42. frogs of Borneo. – The Raffles Bulletin of Zo- Matsui, M. (2005): Taxonomic relationships of ology, 44: 55-574. two species of Rhacophorus from Vietnam Inger, R.F., N. Orlov & I. Darevsky (999): (Amphibia, Rhacophoridae). – Current Her- Frogs of Vietnam: a report on new collections. petology, 24: 9-93. – Fieldiana Zoology, New Series, 92: iii-iv + - Matsui, M. & S. Panha (2006): A new species 46. of Rhacophorus from eastern Thailand (Anu- Inger, R.F., R.B. Stuebing & F.L. Tan (995): New ra: Rhacophoridae). – Zoological Science, 23: species and new records of anurans from Bor- 477-48. neo. – The Raffles Bulletin of Zoology, 43: 5- Myers, C.W. & W.E. Duellman (982): A new 3. species of Hyla from Cerro Colorado, and Inger, R.F., R.B. Stuebing & F.L. Tan (996): other treefrog records and geographical notes Frogs and toads. – pp. 352-367 in Wong, K. from western Panama. – American Museum M. & A. Philipps (eds.): Kinabalu. Summit of Novitates, 2752: -32. Borneo. – Sabah Society, Sabah Parks, Kota Ki- nabalu, Malaysia. Ohler, A. & M. Delorme (2006): Well known does not mean well studied: Morphological Inger, R.F., F.L. Tan & P. Yambun (2000): The and molecular support for existence of sibling frog fauna of three parks in Sabah, Malaysia species in the Javanese gliding frog Rhacopho- - Kinabalu Park, Crocker Range Park, and Ta- rus reinwardtii (Amphibia, Anura). – Comptes wau Hills Park. – Sabah Parks Nature Journal, Rendues Biologies, 329: 86-97. 3: 7-28. Ohler, A., O. Marquis, S. Swan & S. Grosjean Kiew, B.H. (984): A new species of burrowing (2000): Amphibian biodiversity of Hoang Lien frog (Calluella flava sp. nov.) from Borneo. – Nature Reserve (Lao Cai Province, northern Malayan Nature Journal, 37: 63-66. Vietnam) with description of two new species. Malkmus, R. (992): Herpetologische Beobach- – Herpetozoa, 3(/2): 7-87. tungen am Mount Kinabalu, Nord-Borneo. III. – Mitteilungen des Zoologischen Museums Orlov, N.L., A. Lathrop, R.W. Murphy & H.T. Berlin, 68: 0-38. Cuc (200): Frogs of the familiy Rhacophori- dae (Anura: Amphibia) in the northern Hoang Malkmus, R. (993): Zur Zeichnungsvariabilität Lien Mountains (Mount Fan Si Pan, Sa Pa Dis- und zum Totstellverhalten des Ruderfrosches trict, Lao Cai Province), Vietnam. – Russian Rhacophorus angulirostris Mocquard, 890. Journal of Herpetology, 8: 7-44. – Sauria, 5(3): 35-38. Sanchez-Herraiz, M.J., R. Marquez, L.J. Bar- Malkmus, R. (995): Kopfseitenflecken bei bei- badillo & J. Bosch (995): Mating calls of den Geschlechtern von Rhacophorus anguli- three species of anurans from Borneo. – Her- rostris Ahl, 927 (Amphibia: Rhacophoridae). petological Journal, 5: 293-297. – Salamandra, 3: 245-246. Malkmus, R. (2002): Die Amphibien des Mulu- Smith, M.A. (930): The Reptilia and Amphibia of Nationalparks in Sarawak/Malaysia. – Natur the Malay Peninsular. – Bulletin of the Raffles und Museum, 32: 93-05. Museum, 3: -49. Malkmus, R., U. Manthey, G. Vogel, P. Hoff- Stuebing, R.B. & A. Wong (2000): A new species mann & J. Kosuch (2002): Amphibians & Rep- of frog, Philautus erythrophthalmus (Rhaco- tiles of Mount Kinabalu (North Borneo). – Ko- phoridae) from southwestern Sabah, Malaysia. eltz Scientific Books, Königstein. 424 pp. – The Raffles Bulletin of Zoology, 48: 293-296. Manthey, U. & W. Grossmann (997): Amphibi- Taylor, E.H. (962): The amphibian fauna of en & Reptilien Südostasiens. – Natur und Tier Thailand. – University of Kansas Science Bul- - Verlag, Münster. 52 pp. letin, 43: 265-599.

203 J. Maximilian Dehling

Wilkinson, J.A. & R.C. Drewes (2000): Cha- Rhacophorus belalongensis: Brunei Darussa- racter assessment, genus level boundaries, lam: Kuala Belalong Field Studies Centre, Tem- and phylogenetic analyses of the family Rha- burong District (ZMB 70377 [holotype], 70378, cophoridae: a review and present day status. 70379 [paratypes]; UBD GK 06-22, 06-23, 06-24, – Contemporary Herpetology (2): -24. URL: 06-25, 06-26, 06-57, G07-, 07-2, 07-3, 07-4, 07-5, http://www.cnah.org/ch/ch/2000/2/index.htm 07-6, 07-7, 07-8, 07-9 [paratypes]). (26.08.2007) Rhacophorus cf. bipunctatus: Myanmar: “Bir- Wilkinson, J.A. & D. Rao (2004): Taxonomic sta- ma” (ZMB 575, 70072). “Travancore” (ZMB tus of Rhacophorus taronensis Smith, 940. – 03). Proceedings of the California Academy of Sci- Rhacophorus bimaculatus: Philippines: Agusan ences, 55(25): 45-457. River, Mindanao (ZMB 568 [syntype]); Dapitan, Mindanao (SMF 7053). Wilkinson, J. A., R. C. Drewes & O. C. Tatum Rhacophorus cyanopunctatus: Brunei Darussa- (2002): A molecular phylogenetic analysis of lam: Kuala Belalong Field Studies Centre, Tembu- the family Rhacophoridae with an emphasis rong District (UBD GK06-3). Malaysia: Sarawak: on the Asian and African genera. – Molecular Gunung Mulu National Park, Miri Division (ZMB Phylogenetics and Evolution, 24: 265-273. 70707-70709). Thailand: Khao Sok National Park, Wilkinson, J.A., T. Thin, K.S. Lwin & A.K. Shein Surat Thani (ZMB 57895 [holotype]). (2005): A new species of Rhacophorus (Anu- Rhacophorus dulitensis: Brunei Darussalam: ra: Rhacophoridae) from Myanmar (Burma). Batu Apoi, Temburong District (UBD 550). Ma- – Proceedings of the California Academy of laysia: Sabah: Marak Parak, Kota Merudu District Science, 56(4): 42-52. (SP 634-636, 842-844); Hutan Simpan, Mandamui, Wolf, S. (936): Revision der Untergattung Rha- Pitas Kudat (SP 65-68, 20377); Taman Bukit- cophorus. – Bulletin of the Raffles Museum, 2: bukit Tawau, Tawau District (SP 637, 264). 37-27. Rhacophorus edentulus: Indonesia: Northern Ziegler, T. (2002): Die Amphibien und Reptili- Sulawesi (ZMB 34323). en eines Tieflandfeuchtwald-Schutzgebietes in Rhacophorus everetti: Malaysia: Sabah: Marai Vietnam. – Natur und Tier - Verlag, Münster. Parai (SP 362, 23); Sayap, Kota Belud District (SP 342 pp. 202, 203, 773, 865, 285); Headquarters, Kina- balu Park, Ranau District (SP 2700, 270, 20020, Ziegler, T. & J. Köhler (200): Rhacophorus or- 20322, 20325, 20327, 20362, 242, 245, 2452); lovi sp. n., ein neuer Ruderfrosch aus Vietnam Trusmadi (SP 289, 290). (Amphibia: Anura: Rhacophoridae). – Sauria, Rhacophorus fasciatus: Malaysia: Sarawak: Gu- 23(3): 37-46. nung Mulu National Park, Miri Division (ZMB 7070). Rhacophorus gadingensis: Malaysia: Sarawak: Appendix Gunung Gading National Park, Lundu Divisi- Comparative material examined on (ID[Indraneil Das, field number]-783, -7832 [holotype and paratype, respectively]; compari- Rhacophorus angulirostris: Malaysia: Sabah: Gu- son based on detailed photographs provided by I. nung Trusmadi (SP 2859, 2869, 2884, 2896, 2904, Das). 293), Sungai Silau-Silau, Headquarters, Kinabalu Rhacophorus gauni: Malaysia: Sarawak: Mengi- National Park (ZMB 49022, 70073). ong River, Nanga Tekalit, Third Division (FMNH Rhacophorus appendiculatus: Malaysia: Sabah: 3798, 37983, 37985, 39343, 39344, 39346 [pa- Taman Bukit Tawau, Tawau District (SP 098-0, ratypes], FMNH 45542, 45545, 46269, 95359, 2355, 26064); Linumunsut Lake, Maliau, (SP 287); 95445-95449); Nanga Tekalit, Seventh Division Lower Segama, Lahad Datu District (SP 20370- (FMNH 22743, 22744, 22746-22748); Gunung 20372, 20374, 20375). Philippines: Culion, Calami- Mulu National Park, Miri Division (ZMB 707- an Group (SMF 6984); Northeast Mindanao (SMF 7077). Sabah: Sungai Kilampun, Purulon Camp, 6985, 6986); no locality (ZMB 5464, 7007). Crocker Range National Park (FMNH 239235; SP Rhacophorus baluensis: Malaysia: Sabah: Head- 272, 276); Mendolong Camp, Sipitang District quarters, Taman Kinabalu, Ranau District (SP 24, (FMNH 235045, 239236, 239240, 242922, 242923, 29); Restaurant Bayu, near Kinabalu Park, Ra- 24294, 242926; SP 278, 279); Poring Station, Mt. nau District (SP 2802). Kinabalu Park, Ranau District (FMNH 248308;

204 A new Rhacophorus from Borneo

SP 257, 805); Tawau Hills Park, Tawau District Rhacophorus pardalis: Brunei Darussalam: Ku- (FMNH 248924, 248925, 249833-249836; SP 645, ala Belalong Field Studies Centre, Temburong Dis- 646, 072); Marak Parak, Kota Marudu District trict (UBD GK06-07); without locality (UBD 7). (FMNH 235747); Rangkam Kimanis, Pantod Be- Malaysia: Sabah: Taman Bukit Tawau, Tawau Dis- sar, Tambunan District (FMNH 239233); Danum trict (SP 2723, 26060); Danum Valley Field Centre, Valley Field Centre, Lahad Datu District (FMNH Lahad Datu District (SP 2082); Pulau Tiga Natio- 23062, 23069, 2307, 23073, 23075, 234990, nal Park (SP 640-642, 644, 2778-278); Kg. Tipa- 234994, 2408, 24083, 24085, 24086, 24090, su, Napong , Ranau District (SP 2033); Mongko- 245890, 245892, 245894, 245904, 245905, 245909, po, Ranau District (SP 2986); Sungai Kokoguan, 24590, 24593, 24595, 245920); Sungai Agathis, Marak Parak, Kota Marudu District (SP 353, 2083); Maliau Basin (SP 20244). Mendulong, Sipitang District (SP 97-99, 2084); Rhacophorus georgii: Indonesia: Tanke Solok- Maliau Basin (SP 20255-20257); Sungai Rompon, ko, Mekongga Mountains, Southeast Sulawesi Trusmadi (SP 67); Pulau Jembongan (SP 290); (ZMB 34322). Lower Segama, Lahad Datu District (SP 20378, Rhacophorus harrissoni: Brunei Darussalam: 2038, 20382, 20384, 20389); Hutan Simpan, Men- Batu Apoi, Temburong District (UBD 24). Ma- damai, Pitas Kudat (SP 69-80); no locality laysia: Sabah: Lower Segama, Lahad Datu Dis- (SP 688, 694, 696); PDC Lembak Inbak, Telu- trict (SP 20392-20394); Maliau Basin (SP 20279- pid (SP 2660-266); Malangkap Tomis, Kinabalu 2028). Park, Kota Belud District (SP 20768-2077); Sg. Rhacophorus monticola: Indonesia: Southern Kimanis, Kg. Kindosodon, Tambunan District (SP Sulawesi (SMF 6829). 2634). Philippines: Palawan (SMF 6994); Clave- Rhacophorus nigropalmatus: Brunei Darussa- ria, Northern Luzon (SMF 6995). lam: Batu Apoi, Temburong District (UBD 366). : Borneo (SMF 76372 Malaysia: Sabah: Sungai Stuebing, Trusmadi, Tam- [two specimens]). bunan District (SP 223); Tawau Hills Park, Tawau Rhacophorus rufipes: Brunei Darussalam: Kua- District (SP 286); “primary forest” (SP 20696). la Belalong Field Studies Centre, Temburong Dis- trict (UBD GK06-08).

Manuscript received: 28 March 2008 Author’s address: J. Maximilian Dehling, Department of Ecology and Tropical Biology, Bio- centre, University of Würzburg, Am Hubland, D-97074 Würzburg, Germany, E-Mail: maxdehling@web. de.

205 J. Maximilian Dehling

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