NEW AND POORLY KNOWN PARACHUTING FROGS (RHACOPHORIDAE: RHACOPHORUS) FROM SUMATRA AND JAVA

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1 Herpetological Monographs, 16, 2002, by The Herpetologists League, Inc. NEW AND POORLY KNOWN PARACHUTING FROGS (RHACOPHORIDAE: RHACOPHORUS) FROM SUMATRA AND JAVA MICHAEL B. HARVEY 1,AARON J. PEMBERTON 1, AND ERIC N. SMITH 2 1 Department of Biological Sciences, East Tennessee State University, Johnson City, TN , USA 2 Department of Biology, Box 19498, University of Texas at Arlington, Arlington, TX , USA ABSTRACT: We report on a small collection of parachuting frogs from Sumatra and Java. Three new species are described. Rhacophorus achantharrhena is similar to R. dulitensis and R. prominanus and differs from these species by a suite of characters including morphology of the supratympanic fold, digital webbing, coloration, and morphometrics. These three species are unusual in having white visceral and parietal peritonea. Rhacophorus catamitus is a small species similar to R. angulirostris and differing from this species by having a calcar at the heel and reduced digital webbing. Rhacophorus barisani resembles R. baluensis but differs from this species in color pattern, habitus, webbing of the fingers, and morphology of the dermal appendages. A new specimen of Sumatran R. pardalis is described and compared to the holotype of R. pulchellus. Rhacophorus prominanus is reported from Gunung Rajabasa, Lampung. Two specimens are described and compared to Bornean R. dulitensis and R. prominanus from the Malay Peninsula. Rhacophorus tunkui Kiew is a junior subjective synonym of Rhacophorus prominanus Smith. Finally, we describe new specimens of Rhacophorus margaritifer from Cibodas, Java. Skeletons of the new species and of R. margaritifer are described in detail. Superficial jaw and throat musculature appears to be relatively conservative within the genus. Key words: Rhacophorus achantharrhena new species; Rhacophorus catamitus new species; Rhacophorus barisani new species; Rhacophorus prominanus; Rhacophorus pardalis; Rhacophorus margaritifer; Osteology; Sumatra TREE FROGS of the genus Rhacophorus are sometimes called parachuting frogs or flying frogs, because some species possess extensive digital webbing and use their hands and feet like parachutes when escaping predators (Davis, 1965). The genus has its center of diversity on the landmasses of the Sunda Continental Shelf, with more than half of the 37 recognized species occurring on Malaysia, Sumatra, Borneo, and Java (Frost, 2001; Inger and Tan, 1996, Iskandar and Colijn, 2000). Bornean and Malaysian Rhacophorus have been revised recently (Berry, 1975; Inger, 1966) and are well represented in museums. In contrast, species from Sumatra have rarely been mentioned since the work of Boulenger and Van Kampen. Herpetological research on the island of Sumatra has lagged far behind that on adjacent islands of the Sunda Shelf. Over the past century, ten species of parachuting frogs have been discovered on this island and the small islands off its west coast. Rhacophorus nigropalmatus (as Rhacophorus Reinwardti) was the first member of the genus to be reported (Hubrecht, 1887). This species was later found on Laut Tador (Werner, 1900) and in the vicinity of Tandjong Kassan (Werner, 1900) and Palembang (van Kampen, 1905). Collections in the Mentawai Islands made by Modigliani and Maas between 1880 and 1890 included R. appendiculatus and R. pardalis (Boulenger, 1894; Dring et al., 1989), and these species were later reported from the mainland by Werner (1900) who mistakenly named them as new species: Rhacophorus phyllopygus ( R. appendiculatus, type locality Indragiri) and Rhacophorus pulchellus ( R. pardalis from Djapura). Werner (1900) also thought that the population of R. reinwardtii on Laut Tador in the Batu Islands was distinctive enough to warrant the name Rhacophorus reinwardti var. lateralis. Although van Kampen (1923) later agreed that this population has more extensive pigmentation on its flanks, he did not recognize the population as a species distinct from mainland R. reinwardtii. Incidental to other work, Robinson and 46

2 2002] HERPETOLOGICAL MONOGRAPHS 47 Kloss made a small collection of reptiles and amphibians from two rivers: Sungai Kumbang and Sungai Kring on Gunung Kerinci. From this collection, Boulenger (1920) described Rhacophorus modestus and R. poecilonotus. He referred some of the Kerinci material to Werner s R. phyllopygus; however, the specimens from Kerinci were collected at 1426 m. At least in northwestern Borneo, R. appendiculatus occurs at much lower elevations ( m, Inger and Stuebing, 1992). Van Kampen (1923) reported on material collected by L. Ph. de Bussy, E. D. W. Jacobson, and F. C. van Heurn in Sumatra. From the type locality of R. poecilonotus, he described R. bifasciatus as new and reported new Sumatran locality information for R. dulitensis (from Sungai Kring and S. Kumbang), R. reinwardtii ( Lauttador Laut Tador), and R. nigropalmatus (Laut Tador, Tandjungkasau, Maurolabu!, Benakat). Wolf (1936) relegated many of the Sumatran parachuting frogs to subspecific status of wide ranging species. He accepted van Kampen s conclusions regarding the status of Werner s species, and he made Rhacophorus modestus a subspecies of R. buergeri, and R. poecilonotus and R. angulirostris subspecies of R. schlegelii. He also synonymized R. bifasciatus with R. s. poecilonotus and R. prominanus with R. dulitensis. However, few of Wolf s taxonomic changes gained acceptance, and his methods have been frequently criticized (e.g., Inger, 1966; Inger et al., 1999). In 1966, Inger reviewed Bornean parachuting frogs and compared them to Sumatran specimens of R. angulirostris, R. appendiculatus, R. dulitensis, R. nigropalmatus, and R. pardalis. He also reported a specimen of R. prominanus from Nias Island and predicted that this species also occurs on the mainland. Rhacophorus bimaculatus was reported from Leuser National Park (Malkmus and Kunkel, 1992). Manthey and Steiof (1998) restricted the name R. bimaculatus to a species occurring in the Philippines and proposed the new name Rhacophorus cyanopunctatus for frogs previously referred to R. bimaculatus from Sumatra, Borneo, and the Malay Peninsula. Several species originally described from Sumatra and placed in Rhacophorus were later shown to belong to other genera. The holotype of Rhacophorus colletti (now in Polypedates) was collected at Langkat, northeastern Sumatra (Boulenger, 1890). Rhacophorus anodon was described in 1907 (van Kampen, 1907) from near Ajer Mantjur, Kajau Tanam, but was later placed in Philautus (van Kampen, 1923) and shown to be a synonym of Nyctixalus pictus. Ahl s (1927) Rhacophorus kampeni from Sumatra was shown to be a synonym of Polypedates leucomystax (Wolf, 1936). The content and monophyly of Rhacophorus has long been in question. Dubois (1986) considered Polypedates to be a synonym of Rhacophorus and resurrected Leptomantis Peters as a subgenus containing R. bimaculatus, R. gauni, and R. oxycephalus ( Buergeria oxycephalus). He also recognized ten groups of Rhacophorus but did not define these groups with characters. Later (Dubois, 1992), he transferred R. angulirostris to Leptomantis, citing Inger and Tan s (1990) description of the species s tadpole. More recently, Iskandar and Colijn (2000) elevated Leptomantis to genus provisionally considered as valid and expanded it to include R. appendiculatus and species of Philautus in Dubois s (1986) subgenus Gorhixalus (the Philautus hosii group of Dring, 1987). In defense of their action, they listed three characters that they believe define Leptomantis (1) species of small to medium size, (2) occurrence in lowlands, (3) eggs laid in a foam nest and known tadpoles with sucker-like mouths adapted to medium current. Without explanation, these same authors referred R. kajau, R. bisacculus, R. everetti, and R. exechopygus to Rhacophorus sensu stricto, even though these taxa share derived characters (i.e., synapomorphies) with R. appendiculatus (Dring, 1983; Inger et al., 1999; Liem, 1970). Channing s (1989) reanalysis of Liem s (1970) data showed that Polypedates and Rhacophorus are not sister groups. However, a more recent reanalysis of Liem s

3 48 HERPETOLOGICAL MONOGRAPHS [No. 16 data (Wilkinson and Drewes, 2000) found that these genera form a clade exclusive of other rhacophorid genera. We recognize these genera as distinct and defined by characteristics presented in Liem s (1970) table 5. Although Rhacophorus may not be monophyletic (Emerson et al., 2000), we do not believe that Leptomantis is a natural group and believe that even Dubois s more conservative arrangement is premature at best and misleading at worst. Herein, species of Iskandar and Colijn s Leptomantis hosii group are considered to be members of Philautus, and the other species of Leptomantis are considered to be in Rhacophorus. We do not attempt to assign species to Dubois s (1986) ten groups, because he does not define these groups using characters. Our new species are referred to Rhacophorus because they possess the characteristics diagnostic of that genus (Brown and Alcala, 1994, provide the most recent diagnosis of Rhacophorus). While conducting fieldwork in Sumatra in 1996, Harvey and Smith made a small collection of parachuting frogs. The collection included at least four species hitherto unreported from Sumatra. Herein we report on this collection and take this opportunity to redescribe Rhacophorus margaritifer, a poorly known species from Java. MATERIAL AND METHODS Harvey and Smith collected near Bogor in West Java ( Jawa Barat) between May 1996 and during the following dates and times in Sumatra: Bukit Barisan Mountain Range of Bengkulu, 29 May June 1996; Bukit Barisan Mountain Range of Jambi, 6 June June 1996; Gunung Rajabasa, Lampung, 15 June Rains fell almost daily during this period, though often for short periods of time. With a dial caliper under a dissecting scope and to the nearest 0.01 mm, we measured snout vent length (SVL), head length (HL, from the rictus to the tip of the snout), head width (HW, at the level of the rictus), eye diameter (ED), eye-nostril distance (EN, from the anterior border of the ocular aperture to the center of the nostril), distance from the nostril to the tip of the snout (NS), length of the tympanum and its annulus (TympL), eyelid width (EW, maximum distance from the orbital margin to the edge of the eyelid), interorbital distance (IOD, shortest distance between the orbits), internarial distance (IND, distance between the upper edge of each nostril), hand length (HL, from the proximal edge of the palmar tubercle or tubercles to the distal margin of the third finger), foot length (FL, from the proximal margin of the inner metatarsal tubercle to the distal margin of the fourth toe), length of the shank (TL), and the width of the digital tips of the first finger (DF1W), third finger (DF3W), and fourth toe (DT4W). In parentheses, means standard deviation follow ranges of morphometric characters. A slash mark (e.g., 6/7 teeth) separates meristic data from opposite sides of the same specimen, whereas a dash is used to report ranges from more than one specimen or among structures. Structures that were not easily seen (muscle fibers, nonkeratinized nuptial excrescences, characters inside the mouth, and supernumerary tubercles on the hands of some species) were stained reversibly with methylene blue in 70% ethanol. We cleared and double stained skeletons using the procedure of Taylor and van Dyke (1985). Our notation for webbing formulae is that of Savage and Heyer (1967) as modified by Myers and Duellman (1982). Our terminology for the osteological descriptions is that of Trueb (1970, 1993) and Duellman and Trueb (1986). Although we identified our new species on the basis of discrete morphological characters, we applied principal component analysis to morphometric data for two groups of superficially similar species: (1) Rhacophorus barisani and R. baluensis and (2) R. achantharrhena, Bornean R. dulitensis, and Sumatran R. prominanus. Rhacophorus are sexually dimorphic and only adult males (all specimens had nuptial excrescences) were used. Thirteen mensural characters were used to derive components based on 16 (7 R. baluensis and 9 R. barisani) and 19 (9 R. achantharrhena, 8 R. dulitensis, and 2 R. promianus) ob-

4 2002] HERPETOLOGICAL MONOGRAPHS 49 FIG. 1. Rhacophorus achantharrhena (UTACV 53962, SVL mm), an adult female from Bukit Kaba. servations. Our sample sizes are not very large, however we use this method only as an exploratory tool and not for variable (i.e., character) reduction for subsequent analysis. Specifically, the method is used to visualize morphological differentiation among species and to identify mensural characters that may be diagnostic. In contrast to statistical tests such as MANOVA, specimens are not assigned to taxonomic groups a priori in principal component analysis. We examined both the raw data and logtransformed data through normal probability plots and simple distribution graphs for each variable per taxon and for pooled taxa. At the univariate level, we detected no deviations from normality and multivariate normality was assumed. We believe this assumption to be justified because principal component analysis is robust even to relatively extreme departures from normality (Kleinbaum et al., 1988). Because of the specific goals, we used raw data in the principal component analysis: variables were not transformed and components were not rotated. The Kaiser criterion (Statistica, 1991) was applied for selecting the number of principal components: only components with eigenvalues greater than 1 were retained. This criterion was found to be more conservative than the commonly used scree test (Hair et al., 1987). Using the extracted components as axes, we graphed individual specimens to assess whether detectable structure in the data supported our taxonomic assignments. We performed all statistical calculations with the Microsoft Excel 2001 and Statistica (Statsoft, 1994) software packages. SYSTEMATICS Rhacophorus achantharrhena sp. nov. Fig. 1 Holotype. An adult male, UTACV (collector s tag M. B. Harvey 5390), collected 31 May 1996 by Michael B. Har-

5 50 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 2. Distributions of six species of parachuting frogs in Western Indonesia. vey and Eric N. Smith on the western slope of Bukit Kaba, Bengkulu, Sumatra, Indonesia, S, E, 1415 m (Fig. 2). Paratypes (9). Seven adult males, UTACV (M. B. Harvey 5389, M. B. Harvey , ENS ), and one adult female, UTACV (M. B. Harvey 5388) collected with the holotype and one adult male, UTACV (M. B. Harvey 5454), from the southeastern slope of Gunung Dempo, Sumatera Selatan, Sumatra, Indonesia, S, E, m collected 3 June Diagnosis. The new species may be distinguished from all its congeners by the following combination of characters: (1) the SVL of males reaches mm and that of females reaches mm. (2) The snout of males slopes in lateral view and is acuminate in dorsal view; the tip is weakly spatulate. The snout of females is more rounded and not weakly spatulate. (3) The nares are closer to the tip of the snout than to the eye. (4) Webbing of the hand is moderate: web is evident though fails to reach the proximal subarticular tubercles between fingers 1 and 2; it reaches the penultimate phalanx along the postaxial edge of finger 2 and preaxial edge of finger 4 but does not extend beyond the distal subarticular tubercle along the preaxial edge of finger 3. (5) Thick, smooth dermal flaps extend along the postaxial margins of the forearm and tarsus and continue onto the elbow and heel where these flaps are widest. (6) Calcars are absent. (7) The dorsum is smooth in females and finely spiculate in males. (8) The supratympanic fold is thick and conspicuous. (9) The vent is projecting and a free supracloacal fold is present above it. Noticeably enlarged tubercles are absent below the vent. (10) The dorsum is green in life (yellow in preservative) with black and white spots; cross bands are absent from the limbs. In life, webbing between toes 3 5 is red; the re-

6 2002] HERPETOLOGICAL MONOGRAPHS 51 TABLE 1. Factor loadings (not rotated) of the first two principal components obtained through a principal component analysis of thirteen morphometric characters of species allied to Rhacophorus achantharrhena. Heavily loading variables with relatively greater discriminating power are in bold for emphasis (see text for discussion). Variable Snout-vent length Head length Head width Eye diameter Eye-nostril Nostril-tip of snout Length of tympanum Interorbital distance Internarial distance Length of shank Disc of third finger Disc of first finger Disc of fourth toe Eigenvalue % Variance Principal component Principal component maining webbing is immaculate green. (11) Ventral to the liver, the parietal peritoneum is white. The visceral peritoneum of the digestive tract posterior to the liver and the testes is also white. Comparisons. The new species is most likely to be confused with two other medium-sized green Rhacophorus with red pedal webbing and white peritonea. Rhacophorus dulitensis and R. prominanus both differ from R. achantharrhena (characteristics in parentheses) by having low indistinct supratympanic folds that do not overlap the tympanum or its annulus (a heavy and distinct supratympanic fold overlaps the upper edge of the tympanum and its annulus), sharp dark lines below their canthi rostralis (dense melanophores present but usually not forming a sharp line below), and webbing extending beyond the subarticular tubercles of fingers 1 and 2 (webbing basal between fingers 1 and 2) and reaching or approximating the disks on finger 3 (webbing not extending beyond the distal subarticular tubercle on the preaxial side of finger 3). All three species have a similar distribution of concentrated melanophores at the knees, along the canthal and rostral ridges, and in the skin of the tympanum, however the dorsal FIG. 3. Principal component scores for Bornean Rhacophorus dulitensis and Sumatran R. achantharrhena and R. prominanus. Elipses represent 95% confidence limits. spots of Rhacophorus prominanus and R. dulitensis are noticeably smaller than the dorsal blotches of the new species. Our principal component analysis revealed morphometric differences among these superficially similar species. The all positive factor loadings (Table 1) of principal component 1 indicate that it represents differences in overall size: Rhacophorus prominanus is larger than both the new species and R. dulitensis (Fig. 3). Eye diameter, internarial distance, and diameter of the disk of the fourth toe heavily load principal component 2. This weighted linear combination of the variables accounts for the largest total variation in the data among males of the new species and male R. dulitensis (Table 1; Fig. 3): eye diameter and internarial distance are relatively larger in R. dulitensis compared to R. achantharrhena, whereas width of the disk of the fourth toe is relatively smaller. Description of the holotype. The holotype is an adult male with a relatively small head 108% as wide as long and accounting for 29% of SVL. In dorsal view, the snout is subacuminate, and it slopes gently to a weakly spatulate rostral tip in profile (Fig. 4). The canthus is weakly curved and rounded, and the nostril is slightly swollen. A rounded ridge slopes from the nostril almost to the tip of the snout; the rostral area between the ridges is slightly depressed. The weakly concave lore slopes to the mouth, and the lips are not flared. Eye-nostril distance accounts for 53% of the distance from the anterior

7 52 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 4. Head shape of four species of Rhacophorus: A.Rhacophorus barisani, holotype, UTACV 53971; B. Rhacophorus margaritifer, UTACV 54009; C. Rhacophorus catamitus, holotype, UTACV D. Rhacophorus achantharrhena, holotype, UTACV Each specimen is a male. border of the eye to the tip of the snout, and the nostril is 88% closer to the tip of the snout than to the eye. The eye is directed anterolaterally, and the upper eyelid is 72% as wide as the interorbital region. The tympanum and its annulus are visible, and their length is 71% of the length of the eye. A thick supratympanic fold conceals the dorsoposterior edge of the tympanum and extends onto the flanks to the posterior edge of the scapula. Except for a few concealed surfaces such as the posterior thigh and groin, the body surfaces are finely shagreened with tiny pointed spicules. Spicules are also present on the palmar and plantar surfaces and across the venter. Apart from tiny spicules, the dorsum is smooth and the venter and flanks including the limbs and gular region are areolate. The rictal region lacks discrete tubercles and is areolate. The ventral abdominal surface and the region below the thighs is more coarsely areolate than the gular region. The vent is protuberant. A thick supracloacal fold is free and notched medially; it overhangs the vent, but does not cover it completely. This species lacks a discoidal fold and noticeably enlarged tubercles below the vent. The arms are short and slender; the girth of the upper arm is about two-thirds that of the forearm. The hand is relatively large being 72% as long as the foot. Webbing of the hands is moderate, granular, and somewhat thick (Fig. 5). Webbing between fingers 1 and 2 is evident, though basal, and the formula for the remaining digits is II1 2.25III2 1.25IV. When adpressed, length of finger The digital tip of the first finger is round, and the digital tips of the remaining fingers are subtruncate. Digital pads on the hands and feet are well developed, oval, and surrounded by marginal grooves. Ex-

8 2002] HERPETOLOGICAL MONOGRAPHS 53 FIG. 5. Hand and foot of Rhacophorus achantharrhena, holotype, UTACV cept for a slight depression where the distal phalanges bifurcate, the dorsal surfaces of the digital tips are unmodified. Width of the digital tip of the first finger is 63% and that of the fourth toe is 89% of the width of the digital tip of the third finger. Free portions of the fingers bear moderate keels. A thick keel about one-fourth as wide as the digit extends along the postaxial margin of the fourth finger and is continuous with the ulnar fold. Distal subarticular tubercles of the hand are large and round in frontal as well as sagittal section. On fingers 3 and 4, the proximal subarticular tubercles are tiny, oval, and inconspicuous, being only about one-fourth the width of the digit. They occupy depressed areas of these fingers below the articulation points of the metacarpals and proximal phalanges. The holotype lacks supernumerary tubercles, and the palmar tubercle is low, inconspicuous, and bifid. The thenar tubercle is oval and low; its greatest diameter is about as wide as the digital pad of the third finger. The ventral surface of the arm lacks enlarged tubercles. The ulnar fold is smooth, thick, and about as wide as the third digit. It extends around the elbow to terminate on the distal upper arm. A patch of nonkeratinized and granular nuptial excrescences occupies most of the preaxial margin of the thenar tubercle and extends distally to the level of the subarticular tubercle. The excrescences extend dorsally across most of the skin covering the metacarpal. Nuptial excrescences are not present on any of the other digits. The hind limbs are moderately long and slender; the shank is 47% as long as SVL, and the foot is 87% as long as the shank. As for the hands, webbing of the feet is granular. The pedal webbing formula is I1 1.5II1 2 - III1 2IV2 1 V. When adpressed, the length of toe A thick keel along toe 5 is narrower than the digit and extends onto the tarsus to the heel as a smooth fold. As for the hand, proximal subarticular tubercles are small and inconspicuous on the outer digits. One tubercle on toes 1, 2, and 5 and two tubercles on toes 3 and 4 are large and round in sagittal and frontal section. Supernumerary tubercles and an outer metatarsal tubercle are absent. The inner metatarsal tubercle is flat, oval, and about as wide as the disk on the fourth toe. A transverse fold wider than the digits and continuous with the tarsal fold is present on the heel. Low, oval dentigerous process of the vomer bear 5/6 rounded teeth and are directed posteromedially at about 10 to the transverse plane. At the midline, the processes are separated by a space equal to two times their width, and they are separated from the choanae by spaces equal to about one-half their width. The choanae are small, subcircular, and only partially visible at the edge of the lingual shelves of the maxillae when the specimen is on its back. The tongue is subtriangular, free for just less than one-half its length, and deeply bifurcate posteriorly. The medial notch extends about one-fourth the length of the tongue, and the tongue lacks the median

9 54 HERPETOLOGICAL MONOGRAPHS [No. 16 lingual process (Grant et al., 1997) characteristic of many ranoid frogs. The paired vocal apertures are oval holes situated posterolaterally. Each is about as large as the disk of the first finger. Measurements (in mm): SVL 37.74, HL 10.88, HW 11.80, ED 3.78, EN 2.84, NS 2.49, TympL 2.68, EW 3.04, IOD 4.20, IND 3.20, HL 11.00, FL 15.79, TL 17.56, DF3W 2.03, DF1W 1.28, DT4W Coloration in preservative: The green color of the body has faded to a dull yellow. Clusters of melanophores form blotches across the dorsal body surfaces. These blotches are densest at the knee and along the canthal and rostral ridges. The head and anterior portion of the body is more heavily pigmented than the dorsal pelvic region. The skin of the tympanum bears dense melanophores and stands out against the mostly amelanic skin of the rictal region. Scattered across the dorsum are a few clusters of white chromatophores. Similar white blotches are heavy on the flanks, venter, and rictal area. The free margins of the ulnar, tarsal, and supracloacal flaps are white. The white pigment of the dermis (see internal anatomy below) is visible as a patch extending from the level of the coracoids to the posterior margin of the liver. A ventral blood vessel is visible as a charcoal line just under the skin on the posterior one-half of the venter. It disappears at the level of the pubis. Posterior to the ventral patch of white dermis, some of this pigment extends for a short distance along either side of the ventral vessel. Color in life (based on field notes taken by M. B. Harvey): The dorsum of the holotype was bright green with reddish spots. Dorsal surfaces of the digits, dermal appendages on the limbs and above the vent, and the lower lip were yellowish white. The venter and gular region were bright yellow. Webbing between toes 3 5 was red, whereas the web between the remaining toes and on the hands was green. Except for scattered white blotches, the flanks were translucent above the white parietal peritoneum. The ciliary body of the eye was yellow and the edges of the iris silvery white. A dull red diamond occupying most of the iris surrounded the pupil. The palpebrum was clear except for a narrow band of gray pigment along its distal edge. The conjunctiva was yellow. Variation. Coloration of the paratypes is very similar to that of the holotype. The dorsal spotting is lighter in some specimens (notably UTACV 53968), but some large spots are present in all, and dense clusters of melanophores are always present on the knees, along the canthal and rostral ridges, and in the skin over the tympanum. Dorsally, melanophores are heaviest anteriorly in all specimens. Extent of the white pigmentation in the dermis varies among the type series. At one extreme, the white dermal pigment of UTACV is a midventral patch posterior to the pectoral girdle, not reaching the flanks, and not covering the lateral margins of the liver. At the other extreme, the white pigment of UTACV extends to the level of the clavicles, about half way up the flanks, and posteriorly in the skin covering the middle of the small intestine. In all specimens, a narrow band of white pigment extends along either side of the ventral blood vessel. Prominent, round metacarpal supernumerary tubercles are present in most of the paratypes. They are variable in number, even in the same specimen, but one or two in a row may be present on any of the digits. Supernumerary tubercles on the fourth finger are always, and those on the third usually, two times as large as the proximal subarticular tubercles of these digits. Palmar tubercles are not well defined in any of the paratypes. Variation in webbing among the male specimens of the type series is summarized by II( ) ( )III( ) ( )IV for the hand and I(1 1.5) ( )II(1 1.25) 2III(1 1.5) (2-2.75)IV2 (1 1.25)V for the foot. UTACV has a small circular patch of nuptial excrescences at the level of the phalangeal-metacarpal articulation of the second finger. White glandular tissue is visible below the skin at this location in the other specimens, but they lack nuptial excrescences on the second finger. Among the male type specimens (n

10 2002] HERPETOLOGICAL MONOGRAPHS 55 9), SVLs are ; HL is % ( ) of SVL; HW is % ( ) of HL; EN is % ( ) of snout length; distance from the nostril to the tip of the snout is % ( ) of EN; EW is % ( ) of IOD; IND is % ( ) of IOD; TympL is % ( ) of ED; HL is % ( ) of FL; FL is % ( ) of the TL; TL is % ( ) of SVL; width of first finger is % ( ) and that of the fourth toe is % ( ) of the width of the third finger; and width of the third finger is % ( ) of TympL. Sexual dimorphism. A single female of Rhacophorus achantharrhena is larger than the males (UTACV is mm compared to mm for the male specimens). With few exceptions, morphometric ratios for the female fall within ranges for males. Her head is relatively shorter and wider (HL is 27.7% of SVL; HW is 123.2% of HL); her snout is shorter (EN is 55.8% of snout length); and her nostril is relatively closer to the tip of the snout (the distance from the nostril to the tip of the snout is 79.3% of EN). The female s snout appears more rounded in profile, and the weakly spatulate tip is absent. All body surfaces of the female are smooth or areolate and are devoid of spicules. Webbing is only slightly more extensive; it reaches the base of the disks along the postaxial surface of finger 2 and toes 2 and 3 (complete webbing formulae are II1 2.25III IV for the hand and I II1 2 III1 2IV2 1 V for the foot). No differences in color pattern between the male and female paratypes are apparent. Internal anatomy. The muscles, viscera, and bones were examined for UTACV 53967, an adult male. Extensive patches of white pigment are present in several locations. Deflection of the skin reveals a patch presumably in the basement membrane of the dermis and extending from the level of the coracoids to just anterior of the posterior edge of the liver. A more extensive patch of white pigment is in the parietal peritoneum. It extends from the level of the clavicles to just beyond the trilobed liver and extends into the parietal pericardium to cover the ventral and lateral surfaces of the heart. This patch also extends along either side of the midventral blood vessel visible through the skin. The visceral peritoneum of the lower part of the stomach (pyloric region and lower body of the stomach) not covered by the liver and the intestines is white ventrally and laterally. Dorsally, the visceral peritoneum of these organs is patchily white. The left testis is located slightly more caudad than the right and its visceral peritoneum is entirely white. The visceral peritoneum of the right testis is white except for a small pink portion abutting the liver. The peritoneum of the other organs lacks white pigment. In male Rhacophorus achantharrhena, the Wolffian ducts are highly convoluted with short branches extending to the anterolateral portions of the kidney. A bottleshaped vesicula seminalis (Bhaduri and Mondal, 1965) is absent. Dermal Roofing Bones: The roofing bones of the skull consist of the nasals, sphenethmoid, frontoparietals, and otoccipitals. The halves of the sphenethmoid are fused ventrally and dorsally to completely encircle the anterior end of the cranial cavity (Fig. 6). Dorsally, the sphenethmoid is broad, heavily exostosed, and slightly concave; its antero-lateral portions are overlapped by widely separated, quadrangular nasals. The postero-ventral portion of the sphenethmoid is cartilaginous and surrounds the optic foramen; anteriorly the ventral surface of the sphenethmoid is ossified and heavily exostosed. The maxillary process of the nasal is a broad, short process, which extends postero-laterally to terminate just dorsal to the edge of the pars facialis; the nasal and maxilla are not in contact. The frontoparietals are rectangular, exostosed elements; anteriorly they overlap the sphenethmoid slightly and are separated by a slender fenestra extending approximately ⅓ of their length. Posterior to the fenestra, the frontoparietals become fused to one another and to the heavily exostosed otoccipital. Laterally,

11 56 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 6. Skull and pectoral girdle of Rhacophorus achantharrhena, UTACV Bone is stippled and cartilaginous structures are shaded gray. the frontoparietals form a narrow shelf extending into the orbits to the dorsal edge of the sphenethmoid. Suspensorium: The squamosal is triradiate and oriented anterodorsal to the maxillary arcade at 48 ; its otic ramus is broad and about three-fourths as long as the ventral ramus. The otic ramus overlaps the otic process of the quadrate and directly contacts the otoccipital. The zygomatic ramus is thin and about one-fourth as long as the ventral ramus; it extends to a broad terminus just anterior to the tympanic annulus. The anterior face of the ventral ramus of the squamosal articulates with the posterior face of the quadratojugal. The medial ramus of the pterygoid does not articulates with the otoccipital, but is braced against the skull by broad overlap of the pseudobasal process of the quadrate. The anterior ramus of the pterygoid broadly articulates with the maxilla and terminates at the edge of the planum antorbitale. The quadrate is entirely cartilaginous. Its pterygoid process extends along the ventral surface of the anterior ramus of the pterygoid for its entire length but is not continuous with the planum antorbitale. Maxillary Arcade: The premaxillae are widely separated and bear 10/12 teeth. Their alary processes are directed dorsolaterally and exhibit a slight posterior inclination in profile. The pars palatina of the premaxilla is longest medially and laterally. The maxilla abuts the premaxilla and just overlaps it laterally; its pars facialis is highest posteriorly where it overlaps the planum antorbitale. In this species, an L-shaped notch is not present where the maxilla overlaps the planum antorbitale. Each maxilla bears 65 teeth. The pars palatina of the maxilla is widest anteriorly. The quadratojugal is a small sliver of bone connected to the maxilla by a short ligament.

12 2002] HERPETOLOGICAL MONOGRAPHS 57 Palate: The vomers are widely separated, and their dentigerous processes extend medially to articulate with the sphenethmoid. In this specimen, four teeth are present on the right vomer and teeth are absent from the left. The vomers approach the neopalatines laterally, but do not contact the maxillae or the neopalatines. The anterior process of the vomer invests the postero-lateral edge of the solum nasi. The neopalatine is a long slender bone lying medial to the planum antorbitale, overlapping the sphenethmoid, and bracing against the pars palatina and pars facialis of the maxilla. The cultriform process of the parasphenoid is smooth and rectangular. It is relatively short, extending anteriorly for about two-thirds of the length of the orbit. The alae of the parasphenoid are directed postero-laterally and terminate short of the pseudobasal processes. The postero-medial portion of the parasphenoid is fused to the otoccipital. Nasal Capsules: The cavum principale is open laterally and posteromedially. It is partially enclosed by the tectum nasi dorsally, the oblique cartilage laterally, the alary cartilage anteriorly, a wide septum nasi medially, and the solum nasi ventrally. The nasal, solum nasi, oblique cartilage, and the pars facialis of the maxilla expose a large roughly rectangular fenestra on the side of the skull. This fenestra is positioned such that the vomer is visible in profile. The alary process of the premaxilla is completely ossified and braces the capsule anteriorly. The inferior prenasal cartilage extends from the solum nasi to abut the alary process of the premaxilla dorsal to the pars palatina. The alary cartilage is slightly convex anteriorly and forms the anterior border of the nostril. A short superior prenasal cartilage extends laterally from the alary cartilage and braces against the dorso-posterior face of the alary process of the premaxilla. Structure of the septomaxilla is complex, however this bone can be generally described as U-shaped. The closed, anterior end of the septomaxilla is braced against a dorsally directed lamina inferior. The crista subnasalis extends ventrally from the lamina inferior to brace against the anterior border of the pars facialis and pars palatina of the maxilla. A short-cartilaginous spike projects from the postero-lateral corner of the solum nasi and appears to be an incomplete remnant of the planum terminale. A planum transversale extends beneath the nasal to connect the tectum nasi to the planum antorbitale. The planum antorbitale begins beneath the maxillary process of the nasal and terminates in a broad articulation with the pars palatina of the maxilla. Mandible: Meckel s cartilage lacks ossification with the exception of a small region just anterior to its articular surface and another small area where it articulates with the mentomeckelian bone. The mentomeckelian bones share a synchondrotic connection medially and are fused to the dentaries laterally. Dorso-laterally, the dentary overlaps the anterior third of Meckel s cartilage. Ventro-medially, the angulosplenial overlaps Meckel s cartilage along its entire length; it does not contact the dentary or mentomeckelian bone. Anterior to the articular surface of the mandible, the angulosplenial bears a distinct, rounded prominence. Hyoid: The hyoid bears long slender hyale lacking anterior processes. Anterolateral processes might be easily missed, but are present as minute spikes extending from the hyoid plate. Parahyoid bones are absent. The postero-lateral processes are slender, and their length is approximately half the width of the hyoid plate. The posteromedial processes are narrowly separated and heavily dilated anteriorly. The ends of these processes were damaged during preparation; however, the anterior ends are entirely ossified and are not connected to the hyoid plate by cartilaginous stalks as is the case in some ranoid frogs (Liem, 1970, character 10). Postcranial Axial Skeleton: Rhacophorus achantharrhena possesses eight nonimbricate, procoelus, vertebrae. Neural spines are reduced on presacrals I-III, and absent on all others; the atlantal cotyles are widely separated. Fusion of elements is not apparent, however, the centra and neural arches are exostosed. The transverse processes of presacrals II and IV VIII are ap-

13 58 HERPETOLOGICAL MONOGRAPHS [No. 16 proximately equal in length; those of presacral III are slightly longer. A triangular bony spike projects from the anterior edge near the tip of each transverse process on presacral II. This spike is slightly longer than the transverse process. The transverse processes are dilated distally on presacrals III and IV only. These processes are narrow and peg-like on the more caudal vertebrae and relatively long on presacrals II IV. Cartilaginous tips are present on presacral III only. The shape of these cartilaginous tips is unusual; extending posteriorly and medially, they are fanshaped and nearly twice as long as the adjacent osseous portion of the process. The sacral vertebra possesses an anteriorly directed ridge on the dorsal surface of the neural arch; its diapophyses are moderately dilated and directed postero-laterally. The coccyx bears a bicondylar articulation with the sacral vertebra anteriorly and a small cartilaginous tip posteriorly. A dorsal crest with an anterior prominence is present on the coccyx. This crest becomes reduced as it extends caudally. Vestigial transverse processes of the coccyx are absent. Pelvic Girdle: At the iliosacral articulation an oval-shaped sesamoid bone about one-fourth the length of the diapophysis is present in the transverse ligament. The ligament lacks the groove-like articulation with the distal cartilage found in anurans with moderately expanded sacral diapophyses (articulation type IIB, of Emerson, 1979). The ilial shaft is long and bears a dorsal crest that expands vertically as it extends caudally. The anterior end of the ilial shaft bears a cartilaginous tip. The posterior end bears a small lateral protuberance anterior to its fusion with the ishium. The ilium, ischium, and pubis are fused, and these elements are indistinguishable from one another around the acetabulum. Pectoral Girdle: The epicoracoids are incompletely ossified medially. The clavicle is long, straight, and lies ventral to an almost completely ossified procoracoid; it is not fused to the coracoid. The scapula is bicapitate proximally with an anterior and posterior head. The anterior head is fused to the lateral portion of the clavicle and coracoid. The posterior head of the scapula is fused to the dorso-lateral side of the coracoid. The omosternum is bicapitate posteriorly and bears a large, round, tabular cartilage at its anterior end. The arms of the omosternum are wide, yet narrowly spaced and articulate with processes of the procoracoid. The sternum is a long, bony stylus extending caudally from the girdle. The anterior end is approximately three times the width of its posterior end and articulates with the epicoracoids. The posterior end of the sternum bears a large, round, tabular cartilage that is partially ossified. The suprascapula is a broad, fanshaped element heavily ossified except along its posterior and distal borders. A rounded projection extends from the antero-distal corner of the suprascapula. The cleithrum is subtriangular and longest proximally; it overlaps the anterior border of the suprascapula from its articulation with the scapula to the aforementioned anterior projection. The cleithrum covers approximately the anterior third of the suprascapula. Exostosis is present on the medial coracoids and on the scapula and coracoid where these bones fuse near the glenoid cavity. Hands: The hands of Rhacophorus achantharrhena possess 5 mesopodial elements. Two large elements, presumably the radiale and ulnare, articulate with the radio-ulna. The other mesopodial elements are of uncertain homology (Fabrezi and Alberch, 1996, provide a recent discussion of the difficulty associated with assigning homology to carpal elements) and are located distal to the radiale and ulnare and postaxial to the prepollex. A large cuboidal element lies proximal to metacarpal II IV. Two smaller elements are found between the cuboidal element and the prepollex. The prepollex lies proximal to metacarpal I, and is a broad blade-shaped element with a thin, cartilaginous edge. Metacarpal III bears a large distal prominence. No sesamoid elements are present in the hand. The phalangeal formula of the hand is The terminal phalanges of the hands and feet are Y-shaped. Feet: The feet of this species possess 4 mesopodial elements. The proximal and

14 2002] HERPETOLOGICAL MONOGRAPHS 59 distal epiphyses of the tibiale and fibulare are fused. Two other elements of unknown homology are present. One is horizontally elongated and is associated with metatarsals III V. The remaining element is roughly round and lies in between the elongate element and the prehallux. The prehallux is comprised of two segments, the more proximal of which is cubeshaped and nearly completely ossified; the more distal element is a small triangular piece of cartilage. A small oval sesamoid bone partially fits into a shallow depression ventral to the proximal fusion of the tibiale and fibulare. A similar sesamoid bone lies ventral to the articulation of the femur and tibiafibula. The palmar phalangeal formula is Etymology. The name of the new species is a masculine noun in apposition derived from the Greek nouns akantha meaning thorn or prickle and arrhen meaning male. Males of the new species are covered in tiny spicules, and the new name alludes to this sexually dimorphic character. Natural history. At the type locality, calling males were found between hrs on low vegetation along a rocky stream through secondary growth. The stream lacked current; its water was clear, shallow, and restricted to pools separated by dry patches of streambed. At 2100 hrs, the temperature was 19 C. When a full moon appeared overhead, males stopped calling. At Gunung Dempo, the single male specimen was found at 2115 hrs. (air temperature 18 C) calling from low vegetation above a pool of still water separate from, though adjacent to a large, fast flowing stream. This specimen was found where a road through a tea plantation crosses the stream. The nearest forest was about 1 km higher up the mountain where the stream originated. The lower abdomen of the single female is filled with greatly distended ovaries containing seemingly mature eggs (visible through the skin of the flanks). The yolks are pale yellow. To avoid damaging the specimen, we made no attempt to count the eggs; however, the largest eggs protruding through a small gash on the specimens flank were 1.3 mm in diameter. Rhacophorus barisani sp. nov. (Fig. 7) Holotype. An adult male, UTACV (collector s tag M. B. Harvey 5396), collected 31 May 1996 by Michael B. Harvey and Eric N. Smith on the western slope of Bukit Kaba, Bengkulu, Sumatra, Indonesia, S, E, 1440 m. Paratopotypes (8). Eight adult males, UTACV (collectors tags M. B. Harvey , ENS ) collected with the holotype. Diagnosis. The new species may be distinguished from all its congeners by the following combination of characters: (1) the SVL of males reaches mm (females are unknown); (2) the snout slopes in lateral view, projects substantially beyond the lower jaw, and is pointed in dorsal view; (3) the nares are about equidistant between the eye and the tip of the snout; (4) webbing of hands is moderate: web does not reach the proximal subarticular tubercles between fingers 1 and 2, it does not reach the disk along the postaxial edge of finger 2 nor the distal subarticular tubercle along the preaxial edge of finger 3, and it reaches the disk of digit 4; (5) low dermal ridges or tubercles are present along the ulna and tarsus; (6) a dermal flap with long triangular calcar is present at the tibio-tarsal articulation; (7) the dorsum is diffusely covered in tiny, flat, round tubercles; (8) the supratympanic fold is thick and conspicuous; (9) A low supracloacal ridge is present above the vent; noticeably enlarged tubercles are absent below the vent; (10) the dorsal ground color is brown with inconspicuous cross bands; the flanks, groin, and postaxial skin of the thigh is almost black with lightly pigmented spots (white with a tinge of blue in life); (11) the peritonea are devoid of white patches. Comparisons. Males of the following Sunda Shelf Rhacophorus rarely have SVLs exceeding 40 mm and are considerably smaller than R. barisani: R. angulirostris Ahl, R. appendiculatus (Günther),

15 60 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 7. Rhacophorus barisani (UTACV 53979, SVL mm), an adult male from Bukit Kaba. R. cyanopunctatus Manthey and Steiof, R. everetti Boulenger, R. gauni Inger, R. kajau Dring, R. modestus Boulenger, R. rufipes Inger. Among these, perhaps only R. cyanopunctatus could be confused with R. barisani because coloration and pattern of both species is similar (as Rhacophorus bimaculatus, Inger and Stuebing, 1997, supply a color photo of R. cyanopunctatus in their fig. 103). The new species differs from R. cyanopunctatus (characteristics in parentheses) in having a dermal flap and long triangular calcar at the tibio-tarsal articulation (flap and calcar absent) and an unmarked subocular region (subocular region with prominent cream spot). Among the large to medium sized species, Rhacophorus dulitensis Boulenger, R. nigropalmatus Boulenger, R. prominanus Smith, and R. reinwardtii (Schlegel) are unlikely to be confused with the new species because they are mostly bright green rather than brown, lack a triangular calcar, and have more extensive webbing. Pointed snouts and brown to gray dorsal coloration characterize R. fasciatus, R. harrissoni, and R. robinsoni, however, unlike the new species, webbing reaches the disks of the second finger in each of these taxa and each lacks a calcar. A calcar is present on the heel of R. baluensis and R. poecilonotus. The high, angulate head and sharply pointed snout of the new species readily distinguishes it from R. poecilonotus, a species with a strongly depressed head and a rounded snout. The new species is most similar to R. baluensis. Unlike the new species (characteristics in parentheses), R. baluensis has a free supracloacal flap above the vent (supracloacal ridge present but not free distally) and webbing that reaches the base of the digital pad along the postaxial edge of digit 2 (1.5 phalanges free of web). Rhacophorus baluensis is a more robust frog and has well-developed, smooth flaps along the fourth finger, fifth toe, ulna, and tarsus. In R. barisani, these dermal appendages are narrower, and folds along the ulna and tarsus are crenulated. Finally, the venter of R. baluensis is mostly brown,

16 2002] HERPETOLOGICAL MONOGRAPHS 61 TABLE 2. Factor loadings (not rotated) of the first four principal components obtained through a principal component analysis of thirteen morphometric characters of Rhacophorus baluensis and Rhacophorus barisani. Heavily loading variables with relatively greater discriminating power are in bold for emphasis (see text for discussion). Variable Principal component I Principal component II Principal component III Principal component IV Snout vent length Head length Head width Eye diameter Eye-nostril Nostril-tip of snout Length of tympanum Interorbital distance Internarial distance Length of shank Disc of third finger Disc of first finger Disc of fourth toe Eigenvalue % Variance whereas the venter of the new species is cream with black spots. Our statistical analysis of morphometric data for Rhacophorus barisani and R. baluensis extracted four principal components (Table 2). The new species and R. baluensis differ in their scores for principal component 1 (Fig. 8). Examination of loadings for this component indicates that it is influenced primarily by the larger size of R. baluensis. Combinations of the other principal components did not separate the species and explain intraspecific rather than interspecific variation of the morphometric data set. Description of the holotype. The holotype is an adult male. Its head is 96% as FIG. 8. Principal component scores for Rhacophorus baluensis and R. barisani. Ellipses represent 95% confidence limits. wide as long, and HL accounts for 34% of SVL. The snout slopes in profile (Fig. 4), sharply at first then more gently so that it appears weakly spatulate. In dorsal view, the snout is acuminate, and its anteriormost tip is a low, rounded tubercle at the level of the lips. A low dermal ridge forms the slightly concave canthus and extends anteromedially above the swollen nostril to the tip of the snout. The rostral surface between the dermal ridges is depressed giving the snout an angulate appearance. The weakly concave lores slope to the mouth so that flaring of the lips is noticeable behind the eye only. The nostril is only slightly closer to the tip of the snout than to the eye (the distance from the nostril to the tip of the snout is 98% of EN), eye nostril distance accounts for 50.5% of the distance from the anterior border of the eye to the tip of the snout. The upper eyelid is smooth and about 99% as wide as the IOD. IND is 81% of IOD. The tympanum and its annulus are clearly visible below the skin and their length is 58% of the length of the eye. A thick supratympanic fold covers the dorsal edge of the tympanum and extends onto the flanks to the level the postaxial margin of the scapula. Tiny, flat, round tubercles are scattered across the dorsum. On the body, notice-

17 62 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 9. Hand and foot of Rhacophorus barisani, holotype, UTACV able folds are absent except for a low, transverse, glandular ridge above the vent. Below the vent, the skin is areolate, but noticeably enlarged tubercles are absent. The gular region is wrinkled and weakly areolate. Posterior to the margin of the pectoral girdle the ventral body and femur are coarsely areolate. The arms are short; the girth of the upper arm is about three-fourths that of the forearm. The hands have thick, granular webbing, and the webbing formula of the hand is I II III1.5 2 IV (Fig. 9). The length of the hand is 74% that of the foot. When adpressed, length of digit Marginal grooves completely surround well-developed, ovalshaped digital pads. The digital tips are round, and their dorsal surfaces are unmodified. The width of the digital tip of the third finger is 95% of tympanum length. The width of the digital tip of the first finger is 60% and that of the fourth toe 79% of the width of the digital tip of the third finger. A moderate keel extends along the postaxial edge of digit 4 and becomes narrower and crenulated from the level of the proximal subarticular tubercle to the wrist. The subarticular tubercles are well developed and round in both frontal and sagittal section, and the proximal tubercles on fingers 3 and 4 are much smaller than the other subarticular tubercles. Supernumerary tubercles are smaller, low, and round. They are absent below the phalanges, and they are arrayed in single rows of three (on digits 2 and 3) or four (on digit 4) between the proximal subarticular tubercles and the palmar tubercle. A thenar tubercle is oval, low, and nearly twice as large as the bifid palmar tubercle. A low crenulated ridge extends along the ulnar margin of the forearm and is continuous with a low fold postaxial to and below the elbow. Nonkeratinized and granular nuptial excrescences are restricted to the preaxial and dorsal surfaces of fingers 1 and 2. The nuptial pad is most extensive on digit 1 where it extends almost to the disk. On both fingers, the nuptial excrescences form an elevated circular pad above the phalangeal-metacarpal articulation. The hind limbs are long and slender; tibial length is 45.7% as long as SVL and 109.9% as long as FL. Webbing of the feet resembles that on the hands in being moderately thick and granular. The pedal webbing formula is I1 1.5II1 1.75III1 2IV1.75 1V. When adpressed, the length of toe A prominent keel along the postaxial edge of digit 5 becomes low and thick along the edge of the sole of the foot. A short space separates it from a similar thick and weakly crenulated fold extending almost the entire postaxial length of the tarsus. The subarticular tubercles are round to oval in frontal section. As for the hand, the distal subarticular tubercles are round in sagittal section, but the proximal ones project slightly. Supernumerary tubercles are absent below the phalanges and form single rows medial to the proximal subarticular tubercles. The inner metatarsal tubercle is flat, oval, and

18 2002] HERPETOLOGICAL MONOGRAPHS 63 about as large as a the disk of the fourth toe. The outer metatarsal tubercle is absent. The heel bears a postaxial fold with a long triangular calcar distally and a low rounded tubercle proximally. Elevated dentigerous processes of the vomers bear 8/9 subconical teeth, are about twice as wide as long, and are positioned near and medial to the anterior margins of the choanae. The dentigerous processes are about two times as large as the choanae and are separated medially by a space equal to their longest diameter. Each process is directed posteromedially at about 30(to the transverse plane. The small subcircular choanae are positioned far laterally and are just visible at the edges of the lingual shelves of the maxillae when the specimen is on its back. The tongue is subtriangular, free for about one-third its length, and deeply bifurcate posteriorly. The medial notch extends about one-fourth the length of the tongue, and the tongue lacks the median lingual process (Grant et al., 1997) characteristic of many ranoid frogs. The paired vocal apertures are large gaping slits positioned laterally and extending posteriorly beyond the rictus. Coloration in preservative: The dorsum is golden brown with diffuse darker brown blotches. Few, small black dots are scattered more or less evenly across the dorsal body and limbs. Four short charcoal bands cross the shank and two cross the antebrachium. The supratympanic fold is dark brown and contrasts sharply against the lighter skin of the tympanic region and dorsum. The tympanum and its annulus are the same shade of light brown as the rictal region. The supracloacal ridge is slightly lighter than the adjacent dorsal skin. The gular region and ventral body are cream with diffuse, brown mottling. A bold pattern of charcoal with small to large white spots is present on the flanks; on the pre- and postaxial surfaces of the thigh; on the ventral, pre-, and postaxial surfaces of the shank; along the preaxial margin of the arm; and along the preaxial tarsus and dorsal surface of the foot and proximal toes 1 4. Ventral surfaces of the hands and feet are brown and the webbing is uniformly charcoal. Coloration in life (based on field notes of M. B. Harvey): Following fixation and preservation, the coloration of the holotype has changed little. In life, the flanks were charcoal and spots white with a faint bluish tint. The dermal fringes were slightly lighter than the brown dorsal pattern. The iris was brown with the upper half noticeably lighter than the lower half. The ciliary body was yellow and distinct. The palpebrum was clear with a brown distal edge, and the conjunctiva was grayishblue. Measurements (in mm): SVL 50.34, HL 17.23, HW 16.50, ED 6.52, EN 4.42, NS 4.34, TympL 3.76, EW 5.20, IOD 5.24, HL 15.51, FL 20.92, TL 23.00, DF3W 3.59, DF1W 2.16, DT4W Variation. The dorsal pattern of some paratypes is more defined than that of the holotype. When well defined, an irregular dark brown bar crosses the canthus and another crosses the ocular region. Dark brown blotches are present on the dorsum and are Y-shaped just behind the eyes. Most specimens have black flecks scattered more or less evenly (noticeably heavier on the sides of the body in UTACV 53980) across the dorsal body and limbs. The supratympanic fold is always darker than adjacent skin. On both sides, UTACV has a light gray stripe in front of the eye. The tympanum and its annulus are almost always the same color as skin in the rictal region, the only exception being UTACV where a cream spot covers most of the central portion of the tympanum on both sides. The supracloacal fold is usually white (the holotype and one paratype being the only specimens with light brown supracloacal folds). The venter always bears some mottling but varies from nearly immaculate (UTACV 53976) to moderately pigmented (UTACV 53975). Usually, pigmentation of the venter is heaviest posteriorly. The range of variation for webbing between the fingers is summarized by I(2 2.5) ( )II1.5 (3 2 )III2 (1.5 2)IV, and that of the toes by I(1 1 )

19 64 HERPETOLOGICAL MONOGRAPHS [No. 16 (1 2)II1 (1.5 2)III(1 1.5) 2IV2 (1 1.5)V. Among the type series (n 9), SVLs are ; HL is % ( ) of SVL; HW is % ( ) of HL; EN is % ( ) of snout length; distance from the nostril to the tip of the snout is % ( ) of EN; EW is % ( ) of IOD; IND is % ( ) of IOD; TympL is % ( ) of ED; HL is % ( ) of FL; FL is % ( ) of the TL; the TL is % ( ) of SVL; width of first finger is % ( ) and that of the fourth toe is % ( ) of width of the third finger; and width of the third finger is ( ) of TympL. Internal anatomy. UTACV 53973, an adult male, was dissected and cleared and stained. This species s testes are black and massive, each having slightly greater mass than the liver. The Wolffian ducts are highly convoluted with short processes extending to the kidneys. A bottle-shaped vesicula seminalis (Bhaduri and Mondal, 1965) is absent. A few chalky white fibers are visible on the medial side of the m. rectus abdominis, but the visceral, parietal, and pericardial peritonea are not white. The m. submentalis is araphic and about one-tenth as long as the m. intermandibularis. The anterior-most fibers of the m. intermandibularis are directed anteromedially and ventrally cover the lateral portions of the m. submentalis. However, these anterior fibers do not form a supplementary slip and do not appear to insert onto the fascia covering the submentalis. Rather, the medial ends of the fibers of the m. intermandibularis are connected by an aponeurosis that extends to the mandibular symphysis. Posterior to the m. submentalis, the median aponeurosis is large and oval, occupying approximately one-third of the m. intermandibularis and extending to the m. interhyoideus. Fibers of the interhyoideus insert along a median raphe extending from the terminus of the intermandibular aponeurosis. The interhyoideus has virtually no posterior or anterior development (sensu Tyler, 1971b). A moderate (Tyler, 1971a) post-mandibular septum attaches to the posterior edge of the interhyoideus ( postseptal development is absent). Direct myointegumental contact is absent. The mm. geniohyoidei are contiguous for the anterior one-third of their length and arise dorsal to the m. submentalis and m. intermandibularis (i.e., they do not interrupt anterior development of the m. intermandibularis). The mandibular ramus of the trigeminal nerve is superficial to the m. adductor mandibulae (S condition of Starrett, 1968). Discrete slips (sensu Lynch, 1993) of the m. depressor mandibulae are not present. Rather, this extensive muscle fans across the suprascapular and posttympanic regions with superficial fibers arising from the dorsal fascia, otic ramus of the squamosal, and posterior and ventral margins of the annulus tympanicus. Deep fibers of this muscle arise from the ventral arm of the squamosal. Dermal Roofing Bones: The roofing bones of the skull consist of the nasals, sphenethmoid, frontoparietals, and otoccipitals (Fig. 10). The halves of the sphenethmoid are fused dorsally and ventrally, and encircle the anterior braincase. Dorsally, the sphenethmoid is broad, concave, and heavily exostosed. Ventrally, the sphenethmoid is also heavily exostosed and bears an unusual V-shaped projection from its antero-medial border. A cartilaginous portion of the sphenethmoid almost completely encircles the optic foramen and is partially overlapped by a ventral shelf of the frontoparietal. The nasals are curved and quadrangular; they overlap the antero-lateral borders of the sphenethmoid. Each nasal bears a slender, posteriorly directed maxillary process that contacts the pars facialis of the maxilla. The frontoparietals are rectangular and form a slender medial fenestra. Laterally, the frontoparietals protrude as a slender shelf into the medial region of the orbits and bear antero-lateral flanges overlapping the postero-lateral corners of the sphenethmoid. The prootics and exoccipitals are fused to from a single compound otoccipital, which is overlapped anteriorly by the frontoparietals. A triangular flap of carti-

20 2002] HERPETOLOGICAL MONOGRAPHS 65 FIG. 10. Skull and pectoral girdle of Rhacophorus barisani, UTACV Bone is stippled and cartilaginous structures are shaded gray. lage at the anterior edge of the orbit is continuous with the planum antorbitale in this species Suspensorium: The squamosal is triradiate, with the ventral ramus being posteriorly directed and articulating with the dorsal and posterior edge of the quadratojugal. The ventral ramus is directed anteriorly at 51 to the maxillary arcade. The otic ramus is about two-thirds as long as the ventral ramus and expands as a shelf to overlap the otic process of the quadrate and directly contact the otoccipital. The zygomatic ramus is about one-fourth as long as the ventral ramus and ends bluntly. The medial ramus of the pterygoid invests the ventral and medial surface of the pseudobasal process of the quadrate; this ramus approximates but fails to contact the otoccipital. The anterior ramus of the pterygoid extends to terminate just short of the posterior edge of the planum antorbitale in a broad articulation with the pars palatina of the maxilla. The quadrate is entirely cartilaginous. Its pterygoid process runs along the ventro-lateral surface of the pterygoid s anterior ramus for its entire length and terminates short of the planum antorbitale. Nasal Capsules: With the exception of a fenestra between the oblique cartilage and septum nasi and the narial aperture, the cavum principale is enclosed by the septum nasi medially, the tectum nasi dorsally, the alary cartilage anteriorly, the oblique cartilage postero-laterally, and the solum nasi ventrally. The alary process of the premaxillae articulates with the superior prenasal cartilage, a process of the alary cartilage. The inferior prenasal cartilage extends anteriorly from the solum nasi and braces against the pars palatina of the premaxilla. The septomaxilla is a U-shaped bony element; its closed end abuts a small lamina inferior. The crista subnasalis extends ventrally from the lamina inferior as

21 66 HERPETOLOGICAL MONOGRAPHS [No. 16 a broad plate, and only about two-thirds of this element abuts the anterior end of the pars facialis of the maxilla. The remainder extends anterior-medially to just above the articulation of the maxilla and premaxilla. The solum nasi bears a short postero-lateral projection. A planum transversale extends from the dorsomedial edge of the tectum nasi below the nasal to connect the nasal capsule to the planum antorbitale. The planum antorbitale forms the anterior rim of the orbits and is overlapped by the maxillary process of the nasals and the preorbital process of the maxillae; this element broadly articulates with the maxilla. Maxillary Arcade: The premaxilla abuts the maxilla and is overlapped laterally by a short process of this bone. Its alary process is oriented vertically, and its pars palatina is widest medially and laterally. The pars facialis of the maxillae is well developed; the preorbtial process articulates with the maxillary process of the nasal. The pars palatina of the maxilla is broadest anteriorly and narrows slightly as it approaches the maxilla-pterygoid articulation. Each premaxilla bears 11 teeth and each maxilla bears 58 teeth. In this species, an L-shaped notch is present where the maxilla overlaps the antero-ventral corner of the planum antorbitale. The quadratojugal is a small sliver of bone and bears a ligamentous connection with the maxilla. Palate: The vomers are widely separated; they contact the sphenethmoid medially but not the maxillae or neopalatines. The dentigerous processes of the vomer are well developed and bear 6/7 true teeth. The vomers brace the postero-lateral edge of the solum nasi. The neopalatine is a slender element medial to the planum antorbitale. It shares a broad articulation with the sphenethmoid medially and with the pars palatina of the maxilla laterally. The cultriform process of the parasphenoid narrows anteriorly and extends for about two-thirds of the length of the orbit. The alae of the parasphenoid are directed postero-laterally and closely approximate but do not contact the pseudobasal processes. The postero-medial edge of the parasphenoid is not fused to the otoccipital and forms an acute point. Mandible: Meckel s cartilage lacks ossification with the exception of a small area anterior to its articular surface and a small area where it articulates with the mentomeckelian bone. The ossified mentomeckelian bones bear a medial synchondrotic connection and are fused to the dentaries. The dentary covers the dorso-lateral surface of Meckel s cartilage for its anterior third. The angulosplenial covers the ventromedial surface of Meckel s cartilage for its entire length and bears a flat prominence antero-medial to the articular surface of Meckel s cartilage. The angulosplenial does not contact the mentomeckelian bone or the dentary. Hyoid: The hyale lack anterior processes and contain a few small patches of ossification. The anterolateral processes are short, anteriorly directed spikes. Anterior horns of the hyoid are absent. Anteriorly, the hyoid plate bears a poorly developed center of parahyoid ossification. The postero-lateral processes are short, caudally directed spikes. The long, bony posteromedial processes are more heavily dilated anteriorly where they directly abut the hyoid plate (i.e., they are not connected to the plate by cartilaginous segments). The posterior ends of the postero-medial processes are cartilaginous. Pectoral Girdle: The epicoracoids are incompletely ossified medially. The clavicle lies ventral to a partially ossified procoracoid and is not fused to the coracoids medially. Laterally, the coracoid and clavicle are fused to the anterior head of the bicapitate scapula. The posterior head is fused to the dorso-lateral surface of the coracoid. Exostosis is present at on all three elements around the glenoid fossa and also on the medial expansion of the coracoid. At its posterior end, the omosternum forks moderately to articulate with short processes of the procoracoid. It bears a broad tabular cartilage at its anterior end. The sternum is slightly dilated where it contacts the epicoracoids and bears a wide tabular cartilage with a median notch posteriorly. The suprascapula is broad, fan-shaped, and heavily ossified except for its postero-distal borders. The subtriangular cleithrum is longest proxi-

22 2002] HERPETOLOGICAL MONOGRAPHS 67 mally and covers the anterior edge of the suprascapula; it extends from near the scapular articulation to a rounded anterior projection at the distal corner of the suprascapula. Postcranial Axial Skeleton: Rhacophorus barisani possesses eight nonimbricate, procoelus vertebra. Fusion of elements is absent, the centra and neural arches are exostosed, and the atlantal cotyles are widely separated. The neural arches of presacrals I II bear small neural spines. The transverse processes of presacrals II, III, and VIII are perpendicular to the body axis. All other transverse processes are slightly directed posteriorly. The transverse processes of III and IV are wider, are dilated laterally, and have cartilaginous tips. All other transverse processes are peg-like and approximately equal in length; those on presacrals V VIII also bear cartilaginous tips. The neural arch of the sacral vertebra has a low horizontal ridge; the sacral diapophyses are moderately dilated and have a slight posterior orientation. The coccyx possesses a bicondylar attachment to the sacral vertebra, a small dorsal crest restricted to its anterior half, and a large anterior prominence. The posterior tip of the coccyx is cartilaginous. Vestigial transverse processes of the coccyx are absent. Pelvic Girdle: The iliosacral articulation is Emerson s (1979) type IIB. In this species, sesamoid bones are not present in the transverse ligament. The anterior tips of the ilia are cartilaginous. Each ilial shaft is long and bears a dorsal crest that expands dorsally as it extends caudally. The dorsal rim of the acetabulum exhibits slight lateral expansion and bears no protuberances. The pubis is ossified and indistinguishable from adjacent elements. Hands: The hands of Rhacophorus barisani possess five mesopodial elements. Two mesopodial elements are associated with radioulna and presumably are the radiale preaxially and the ulnare postaxially. Three distal mesopodial elements are of unknown homology. A large element is associated with metacarpals II IV. Two smaller elements are postaxial to the prepollex and associated with metacarpal I. The prepollex is large and wedge-shaped with a cartilaginous edge. The distal end of the third metacarpal bears a large rounded prominence. Metacarpal II also bears a prominent ridge on its distal preaxial side. The terminal phalanges of the hands and feet are Y-shaped. The palmar phalangeal formula for the hand is The hand lacks sesamoid bones. Feet: Three mesopodial elements are present in the feet. The fibulare and tibiale are fused proximally and distally. A cubeshaped element lies postaxial to the prehallux. The fourth element is elongate and associated with metatarsals I III. The prehallux consists of two elements, the more distal almost completely cartilaginous and the more proximal osseous. The phalangeal formula for the feet is A small oval sesamoid bone partially fits into a depression ventral to the proximal fusion of the tibiale and fibulare. This species lacks a sesamoid bone at the articulation of the femur and tibiafibula. Etymology. The new name barisani is a masculine noun in apposition and refers to the Bukit Barisan Range. Occurring at high elevations in primary rainforest, the new species is likely endemic to this mountain range. Natural history. The type series was found along a stream trickling through tall, upper montane rainforest. Where the stream flattened out, none of these frogs were found. Rather, they were found calling from vegetation m above deep pools where the stream flowed down a steep, basaltic slope. Foam nests were found floating on the water s surface. The notes of the laugh-like call are short in duration and repeated numerous times in quick succession. Rhacophorus catamitus sp. nov. (Fig. 11) Holotype. An adult male, UTACV (collector s tag M. B. Harvey 5444), collected 3 June 1996 by Michael B. Harvey and Eric N. Smith on the southeastern slope of Gunung Dempo, Sumatera Selatan, Sumatra, Indonesia, S, 103, 08, 40 E, m.

23 68 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 11. Sexual dimorphism of Rhacophorus catamitus: the specimen at left is the male holotype (UTACV 53981, SVL mm), and the specimen at right is the female (UTACV 54001, SVL mm). Paratypes. Eighteen adult males, UTACV and (M. B. Harvey , , ENS ), collected with the holotype and three adult males, UTACV (ENS ) on the same mountain at m ( S, E) and also collected 3 June Referred specimens. An adult female (UTACV 54001) from the Bukit Barisan range near Kepahiang, Bengkulu, S, E, m, collected on 30 May 1996; an adult male (UTACV 54002) from the western slope of Bukit Kaba, Bengkulu, 1415 m, collected on 31 May 1996; two males (UTACV ) from about 24 km (by road) west of Sungaipenuh, Jambi, 1040 m, collected on 7 June Diagnosis. The new species may be distinguished from all its congeners by the following combination of characters: (1) The SVL of males is mm (female 50.56). (2)In males, the snout slopes sharply to a low, swollen rostral tubercle, then back toward mouth; the snout is subacuminate in dorsal view. (3) The nares are closer to tip of snout than to the eye. (4) Webbing of the hand is reduced: webbing is basal between digits 1 and 2, does not reach the proximal subarticular tubercles between digits 2 and 3, and does not reach the distal subarticular tubercles between digits 3 and 4. (5) A row of tubercles or low crenulated fold extends along the postaxial margin of the antebrachium and tarsus. (6) A short and large, subtriangular tubercle is present at the tibio-tarsal articulation. (7) The dorsal body and limbs are shagreened with tiny, low, rounded tubercles; the head and eyelids are smooth. (8) The supratympanic fold is thick and conspicuous; it overlaps the upper edge of the tympanum and its annulus. (9) The supracloacal ridge low and white; large white tubercles are present below the vent. (10) The dorsal ground coloration is brown with darker cross bands; white spots occasionally are present on flanks and below

24 2002] HERPETOLOGICAL MONOGRAPHS 69 eye. (11) The peritonea are devoid of white patches. Comparisons. The new species (characteristics in parentheses) is most similar to other small species of Rhacophorus. Rhacophorus gauni is most similar to the new species; however, R. gauni has more extensive webbing with the web extending beyond the distal subarticular tubercle on the fourth finger (web does not reach the distal subarticular tubercles), has a subtruncate snout in profile (snout angulate in profile), and usually has conical tubercles on the eyelids (eyelids smooth). Large, cream-edged, dark brown spots on the dorsa of many specimens of R. gauni are not present in the new species. Superficially, the new species resembles R. angulirostris. However, in R. angulirostris, webbing reaches the disks of the outer fingers (reduced webbing not reaching the distal subarticular tubercles of the fingers), and the heel and supracloacal regions are unornamented (heel with large subtriangular tubercle and supracloacal region with a wavy white glandular ridge). Rhacophorus appendiculatus, R. everetti and R. kajau have highly tuberculate dorsa (dorsum shagreened) and other distinctive characters that readily distinguish them from the new species. Rhacophorus cyanopunctatus, R. modestus, and R. rufipes lack a large tubercle (present) at the tibiotarsal articulation, and males of the first two species also have more rounded heads (snout angulate in profile). Description of the holotype. The holotype (Fig. 11) is an adult male. Its relatively large head is 113% as wide as long, and HL accounts for 34% of SVL. In profile (Fig. 4), the snout slopes sharply to a swollen point then back again towards the mouth. In dorsal view, the snout is subacuminate. The canthus is straight and sharp and the nostril swollen. A ridge slopes sharply from the nostril to the tip of the snout; the rostral area between the ridges is slightly depressed. The weakly concave lore slopes to the mouth, and weak flaring of the lips is noticeable behind the eye only. The nostril is 73% closer to the tip of the snout than the eye; EN accounts for 58% of the distance from the anterior border of the eye to the tip of the snout. The eyes are directed anterolaterally, and the upper eyelid is about 97% as wide as the interorbital region. The tympanum and its annulus are visible though indistinct; their length is 41% of the length of the eye. A thick supratympanic fold extends onto the flanks to the scapular region. The head, upper eyelids, and arms are smooth, and the dorsal body and legs are shagreened. Small tubercles are present in the rictal area, and large tubercles are present below the vent. On the body, noticeable folds are absent except for a low, transverse glandular fold above the vent. The throat, chest, and ventral surfaces of the arms and shanks are smooth, whereas the ventral body and thigh are coarsely areolate. A discoidal fold is absent. The arms are short and slender; the girth of the brachium is only slightly narrower than the girth of the antebrachium. Webbing of the hands is reduced, granular, and somewhat thick (Fig. 12). Webbing between fingers 1 and 2 is basal and the formula for the remaining fingers is II2 3.5III IV. The length of the hand is 77% that of the foot. When adpressed, length of digit All digital tips are round and bear well developed pads surrounded by marginal grooves. Except for a shallow depression where the terminal phalanges bifurcate, the dorsal surfaces of the digital tips are unmodified. The width of the digital tip of the first finger is 55% and that of the fourth finger is 83% of the width of the digital tip of the third finger. A feeble keel extends along the postaxial edge of the fourth finger and hand to the distal margin of the outermost palmar tubercle. The subarticular tubercles of the hand are round in frontal and sagittal section. On fingers 3 and 4, the distal subarticular tubercles are 2 3 times as large as the proximal subarticular tubercles and flatter in profile. Supernumerary tubercles are low, round, and smaller than subarticular tubercles. They are absent below the phalanges, and they are arrayed in single rows of two (fingers 2 and 4) or three (finger 3) between the proximal subarticular tubercles and the

25 70 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 12. Hand and foot of Rhacophorus catamitus, holotype, UTACV palmar tubercle. The thenar tubercle is oval, low, and larger than the bifid palmar tubercle. A short space separates the palmar tubercle from a row of round tubercles along the ventro-postaxial margin of the antebrachium. A small oval patch of nonkeratinized and granular nuptial excrescences can barely be distinguished on the preaxial and dorsal surface of the first finger. Nuptial excrescences are absent from digit 2. The hind limbs are long and slender; TL is 53% as long as SVL, and the foot is 85% as long as the shank. Webbing of the feet is thinner and less granular than that on the hands. The pedal webbing formula is I II III1.5 3IV V. Length of toe A low keel along the postaxial edge of digit 5 extends onto the foot as far as the proximal end of the metatarsal. The subarticular tubercles are round to oval in transverse section. As for the hand, the distal subarticular tubercles are largest and flatter in sagittal section. Supernumerary tubercles of the foot are flat, round, and indistinct. Below the phalanges, they are absent and they are arrayed in rows medial to the basal subarticular tubercles. The inner metatarsal tubercle is flat, oval, and about as large as the disk of the fourth toe. An outer metatarsal tubercle is absent. A low crenulated fold extends along the postaxial edge of the tarsus, and the heel bears a large, subtriangular tubercle. A transverse fold is not present on the heel. Low dentigerous processes of the vomers are located near the anterior edges of the choanae and bear 4/4 subconical teeth. The dentigerous processes are about as large as the choanae and are slightly wider than long. At the midline, the processes are separated by a space equal to three times their width. Each process is directed posteromedially at about 15(to the transverse plane. The dentigerous processes and choanae are positioned far laterally. The choanae are subcircular and just visible at the edges of the lingual shelves of the maxillae when the specimen is on its back. The tongue is oval, free for just less than one-half its length, and deeply bifurcate posteriorly. The posterior notch extends about one-third the length of the tongue, and the tongue lacks the median lingual process (Grant et al., 1997) characteristic of many ranoid frogs. The paired vocal apertures are large slits positioned laterally and extending posteriorly beyond the rictus. Measurements (in mm): SVL 31.28, HL 10.67, HW 12.03, ED 4.46, EN 2.69, NS 1.96, TympL 1.84, EW 3.75, IOD 3.86, IND 2.67, HL 10.76, FL 13.92, TL 16.43, DF3W 2.26, DF1W 1.25, DT4W Coloration in preservative: Dorsally, the holotype is brown with irregular darker brown bands on the back. A narrow dark brown stripe crosses the snout about halfway between the eyes and nares. It is followed by a similar stripe across the eyelids and interorbital region and a stripe just behind the eyes. The sides of the face are the same shade of brown as the dorsal surface of the head, and the lips are unpat-

26 2002] HERPETOLOGICAL MONOGRAPHS 71 terned. The canthal and rostral ridges and the rostral tubercle are lighter than the adjacent skin. The supratympanic fold and skin covering the tympanum and its annulus are the same color as the adjacent skin of the rictal region. A large, broken white spot is below and between the eye and tympanum above the mouth. Three small white spots form a row dorsolaterally about halfway between the limbs on one side only. The flanks are cream, immaculate in the axilla and groin, and with small brown blotches between these immaculate regions. The supracloacal ridge bears a broken white line, and two large tubercles below the vent are white. The posterior thigh is light tan with a diffuse sprinkling of brown pigment granules not forming any definable pattern. The limbs are banded: five bands cross the shank and three cross the antebrachium. Webbing and the dorsal surfaces of the hands and feet are brown except for fingers 1 and 2 that are mostly cream. Except for blotches below the mandible, the gular region and chest are immaculate cream. Posteriorly, areolate portions of the ventral body surfaces are cream, and the ventral surface of the thigh is pale yellow; both areas bear numerous small brown blotches. Coloration in life (based on field notes of M. B. Harvey): The white blotches on the dorsum and the tubercles below the vent were cream in life. The venter was cream with brown flecks. The groin and ventral surfaces of the thighs, hands, and feet were orange. The iris was dirty yellow with a tinge of green. The ciliary body was not distinct, being only slightly more yellow than the iris. The palpebrum was clear and lack an obvious darker distal edge. Variation. Among the type series, SVLs are mm; HL is % ( ) of SVL; HW is % ( ) of HL; EN is % ( ) of snout length; distance from the nostril to the tip of the snout is % ( ) of EN; EW is % ( ) of IOD; IND is % ( ) of IOD; TympL is % ( ) of ED; HL is % ( ) of FL; FL is % ( ) of TL; TL is % ( ) of SVL; width of first finger is % ( ) and that of the fourth toe is % ( ) of width of the third finger; and width of the third finger is ( ) of TympL. Webbing of the hands and feet is conservative. For the hand, variation is summarized by II(2 2.5) 3.5III2.75 (2 2.75)IV. Webbing is basal between the first two fingers in all specimens. On the foot, variation is summarized by I1.75 (2 2.5)II( ) (2.75 3)III( ) ( )IV( ) ( )V. In four specimens, single small flat tubercles can barely be discerned near the lateral edge of one (e.g., UTACV 53990) or both (e.g., UTACV 53993) eyelids. Tubercles below and beside the vent number The tarsal ridge is always more or less crenulated, but the ulnar margin may have a low and almost completely smooth ridge, a low crenulated ridge, or a row of discrete tubercles. In some specimens, the ulnar ridge continues around the ventral and postaxial edge of the elbow as a low fold. As for the holotype, the male paratypes have relatively small patches of nuptial excrescences restricted to the dorsal and preaxial skin covering the metacarpal and between the distal edge of the thenar tubercle and the metacarpal-phalangeal articulation. These nuptial pads are difficult to see without application of stain. Sexual dimorphism. With some hesitation (no males were found at the same locality and all other specimens come from higher elevations), we tentatively refer a single female (UTACV 54001) to this species. With a SVL of mm, the female is nearly twice as large as males (Fig. 11). In profile, the female s snout is rounded and lacks the low tubercle of males. Webbing is more extensive and reaches the disks of all the toes, the preaxial side of finger 2, and the postaxial side of finger 4. Webbing of the remaining fingers is also more extensive: I II1 2III1 1IV. The dorsal body and limbs are covered in scattered, flat, round tubercles; the raised, rounded tubercles of males are absent. Internal anatomy. UTACV 53985, an adult male, was dissected and cleared and

27 72 HERPETOLOGICAL MONOGRAPHS [No. 16 stained. A few white fibers are present in the parietal pericardium, but the pericardial, parietal, and visceral peritonea are not uniformly white. The testes of this species are pink and small, the mass of each equaling one of the three lobes of the liver. The Wolffian ducts are only weakly convoluted with few processes connecting their curves to the kidneys. A bottleshaped vesicula seminalis (Bhaduri and Mondal, 1965) is absent. The m. submentalis is araphic and about one-tenth as long as the m. intermandibularis. The anterior-most fibers of the m. intermandibularis are directed anteromedially and ventrally cover the m. submentalis. However, these anterior fibers do not form a supplementary slip and do not appear to insert onto the fascia covering the submentalis. Posterior to the m. submentalis, the median aponeurosis is large and oval, occupying approximately one-half of the m. intermandibularis and extending to the m. interhyoideus. Fibers of the interhyoideus insert along a median raphe extending from the terminus of the intermandibular aponeurosis. The interhyoideus has virtually no posterior or anterior development (sensu Tyler, 1971b). A moderate (Tyler, 1971a) post-mandibular septum attaches to the posterior edge of the interhyoideus ( postseptal development is absent). Direct myointegumental contact is absent. The mm. geniohyoidei are contiguous only where they lie above the m. submentalis. They arise dorsal to the m. submentalis and m. intermandibularis (i.e., they do not interrupt anterior development of the m. intermandibularis). The mandibular ramus of the trigeminal nerve is superficial to the m. adductor mandibulae (S condition of Starrett, 1968). Discrete slips (sensu Lynch, 1993) of the m. depressor mandibulae are not present. Rather, this extensive muscle fans across the suprascapular and posttympanic regions with superficial fibers arising from the dorsal fascia, otic ramus of the squamosal, and posterior and ventral margins of the annulus tympanicus. Deep fibers of this muscle arise from the ventral arm of the squamosal. Dermal Roofing Bones: The roofing bones of the skull consist of the nasals, sphenethmoid, partially fused frontoparietals, and the otoccipitals (Fig. 13). The sphenethmoid encircles the anterior region of the brain case, being fused dorsally and ventrally. The dorsal surface of the sphenethmoid is slightly concave, broad, and heavily exostosed. The anterior region of the sphenethmoid is overlapped dorsally by the nasals, ventrally by the neopalatines and vomers. Small, triangular flaps of cartilage extend from the anterior-lateral edge of the sphenethmoid into the orbits. These flaps of cartilage are not continuous with the planum antorbitale. Ventrally, the sphenethmoid is ossified and heavily exostosed anteriorly; portions of this bone remain cartilaginous adjacent to the optic foramina. The frontoparietals bear a T- shaped fenestra, extending posteriorly for half their length. The horizontal portion of the fenestra forms the boundary between the sphenethmoid and the anterior edge of the frontoparietals. A noticeable swelling extends from the frontoparietal at the dorsoposterior edge of the orbit. The prootics and exoccipitals are fused to form a heavily exostosed otoccipital. Nasal Capsules: The narial apertures are oriented laterally. The cavum principale is incompletely enclosed by the septum nasi medially, a wide tectum nasi dorsally, the oblique cartilage posteriorly and laterally, the cup-shaped alary cartilage anteriorly, and the solum nasi ventrally. The alary process of the premaxilla braces the capsules anteriorly. The postero-dorsal face of the alary process articulates with a tabular superior prenasal cartilage, and its postero-ventral portion articulates with an inferior prenasal cartilage that extends to the pars palatina. The plate-like crista subnasalis extends ventrally to brace the solum nasi against the pars facialis of the maxilla. The planum transversale extends below the nasal from the dorsomedial edge of the tectum nasi, however, unlike in the other species, this process terminates below the nasal at mid-length and is widely separated from the planum antorbitale in Rhacophorus catamitus. The oblique cartilage forms the postero-lateral margin of the nasal aperture and bears a small posterior projec-

28 2002] HERPETOLOGICAL MONOGRAPHS 73 FIG. 13. Skull and pectoral girdle of Rhacophorus catamitus, UTACV Bone is stippled and cartilaginous structures are shaded gray. tion. The septomaxilla is U-shaped, and its closed end abuts the lamina inferior. A short process extends from the posterior margin of the solum nasi. Behind the nasal capsules, the vomer is exposed by a large rectangular fenestra and is visible in profile. Suspensorium: The squamosal is triradiate and its ventral process is directed anteriorly at 51 to the maxillary arcade. The zygomatic process of the squamosal is onefourth as long as the ventral ramus and comes to a short, broad termination just anterior to the tympanic annulus. The otic process is two times as long as the zygomatic process and directly contacts the otoccipital. The ventral process is long and narrow and covers the anterior and lateral surfaces of the entirely cartilaginous quadrate. The medial ramus of the pterygoid covers the ventral surface of the pseudobasal process of the quadrate and does not reach the otoccipital. The anterior ramus of the pterygoid is thin and elongate, articulates broadly with the pars palatina of the maxilla. The pterygoid process of the quadrate extends along the ventral surface of the anterior ramus for its entire length and does not fuse to the planum antorbitale anteriorly. The lateral ramus covers the medial surface of the quadrate. Maxillary Arcade: The premaxillae bear 10/11 teeth. Their alary processes are directed vertically with slight posterior inclination. The pars palatina is longest medially and laterally. The anterior edge of the maxilla overlaps the premaxilla laterally. Its pars facialis is relatively tall and the pre-orbital process of the maxilla broadly contacts the maxillary process of nasal. The maxilla bears an L-shaped notch where it overlaps the anterior edge of the planum antorbitale. Each maxilla bears 63 teeth. The quadratojugal is a slender bone with a ligamentous connection to the maxilla.

29 74 HERPETOLOGICAL MONOGRAPHS [No. 16 Mandible: With the exception of a small area anterior to its articular surface and a small area where it contacts the mentomeckelian bone, Meckel s cartilage is completely cartilaginous. The mentomeckelian bones bear a synchondrotic connection medially and articulate with the anterior end of Meckel s cartilage. They are fused to the dentaries. The dentary extends posteriorly to invest Meckel s cartilage anterolaterally. The longer angulosplenial extends posteriorly-medially along Meckel s cartilage and does not articulate with the mentomeckelian bone or the dentary. A rounded prominence is present on the angulosplenial just anterior and medial to the articular surface of Meckel s cartilage. Palate: The vomers are reduced and widely separated; their dentigerous processes overlap the sphenethmoid. The vomers invest in the nasal capsules ventrolaterally and do not contact the premaxillae, maxillae, or neopalatines. Presence of vomerine teeth is polymorphic in this species, and this specimen lacks vomerine teeth. The neopalatines are slender and articulate with the sphenethmoid medially and the maxilla laterally. The cultriform process of the parasphenoid bears a low, broad median keel and extends anteriorly for about three-fourths of the length of the orbit. Posteriorly, the parasphenoid is fused to the otoccipital, and its alae are angled slightly posterior of lateral. The alae approximate the pseudobasal processes but do not contact them. Hyoid: The hyale are slender and recurved. The anterior and postero-lateral processes are approximately equal in length. The postero-lateral process is directed caudally. The postero-medial process is ossified, anteriorly dilated and approximately twice the length of the aforementioned processes; its caudal tips are cartilaginous. Cartilaginous stalks do not connect the postero-medial processes to the hyoid plate. Parahyoid bones are absent. Pectoral Girdle: The medial portions of the epicoracoids are unossified. The clavicle is long, straight, slender and overlaps an ossified procoracoid ventrally. The coracoid and clavicle are fused to the scapula laterally. The omosternum bears a large, dilated tabular cartilage at its anterior end and is broadly forked at its posterior end where it articulates with processes of the procoracoid. The sternum is a slender, bony stylus; anteriorly, it is dilated and articulates with the posterior epicoracoids. The posterior end bears a broad, bicapitate cartilage. The suprascapula is fanshaped and heavily ossified except for its distal and posterior edges. It bears an anterior projection at its distal edge. The cleithrum is subtriangular and overlaps the anterior border of the suprascapula. It extends posteriorly to cover the anterior fourth of the suprascapula and reaches the articular surface of this bone. Pelvic Girdle: The iliosacral articulation is type IIB of Emerson (1979). Sesamoid bones are absent from the transverse ligament of this species. The ilial shaft is long and bears a dorsal crest that expands vertically as it extends caudally. Its anterior tip is cartilaginous and it bears a postero-lateral prominence. A protuberance extends laterally from the dorsal acetabular expansion. The pubis is ossified and indistinguishable from the ischium and ilium. Postcranial Axial Skeleton: This species possesses eight, nonimbricate and procoelus presacral vertebrae. Neural spines are highly reduced on presacrals I III and absent on the remaining vertebrae. Fusion of elements is not observed, the centra and neural arches are exostosed, and the atlantal cotyles are widely separated. The transverse processes of presacrals III and IV are slightly expanded with tabular cartilaginous tips, and the process of the third presacral is slightly wider than the processes of the other vertebrae. The remaining transverse processes are peg-like with small cartilaginous tips. The neural arch of the sacral vertebra has a low horizontal ridge. The sacral diapophyses are slightly larger than the transverse processes, slightly expanded, and directed posteriorly. The coccyx bears a bicondylar articulation with the sacral vertebra and no vestigial processes. The coccyx bears a cartilaginous tip posteriorly and prominent ridge dorsally. Hands: Rhacophorus catamitus possesses five mesopodial elements. The radiale

30 2002] HERPETOLOGICAL MONOGRAPHS 75 and ulnare are associated with the radioulna. The other 3 elements are of uncertain homology and will be described by location. Two of these elements are small and postaxial to the prepollex. A larger element is proximal to metacarpals I III. The prepollex consists of 2 elements, which are ossified with the exception of a cartilaginous edge on the more distal element. Metacarpal III bears a large crest along the preaxial margin of its distal end. Metacarpal II bears a similar though smaller crest. The terminal phalanges of the hands and feet are Y-shaped. The phalangeal formula of the hand is Sesamoid bones are not present in the hand. Feet: Four mesopodial elements are present in the feet. The other elements are of uncertain homology. The fibulare and tibale are fused at their proximal and distal ends. A large elongate element is proximally associated with metatarsals I III. A smaller element is located between the prehallux and the aforementioned larger element. The prehallux consists of 2 components, with the more distal being cartilaginous and the proximal element mostly ossified except for a cartilaginous edge. A small oval sesamoid bone partially fills the depression ventral to the proximal fusion of the tibiale and fibulare. A similar sesamoid bone is present ventral to the articulation of the tibiafibula and femur. Etymology. The specific epithet catamitus is a noun in apposition derived from the Latin name for the cup bearer of the gods in Greco-Roman mythology. Catamitus was a boy kept forever young by Jupiter. Although male Rhacophorus are usually smaller than females of the same species, the difference in size is rarely as dramatic as in R. catamitus. In addition, males of this species have much smaller nuptial pads and testes than congeners examined by us; the Wolffian ducts are only weakly convoluted. Natural history. Rhacophorus catamitus occurs in forest of the Bukit Barisan range. All but the female specimen were collected between 1040 and 1695 m. At the type locality and at Bukit Kaba, males called from low vegetation near streams. FIG. 14. Rhacophorus margaritifer (UTACV 54005, SVL mm), an adult female from Gunung Gede, Java. The single female was inactive atop a leaf about 2 m above a small stream through forest between m. Rhacophorus margaritifer (Schlegel) Fig. 14 Hyla margaritifera Schlegel, 1837:107. Syntypes RMNH; type locality Java. Rhacophorus margaritiferus (Schlegel): Tschudi, 1838:75. Rhacophorus javanus Boettger, 1893:338. Holotype SMF 6982; type locality Vulkan Tjisurupan, West-Java, Indonesia. Rhacophorus javanicus Boettger: Boulenger, 1894:35. (Misspelling). Polypedates javanus (Boettger): Van Kampen, 1923:254. Rhacophorus barbouri Ahl, 1927:45. Holotype ZMB; type locality Westjava, Indonesia. Rhacophorus (Rhacophorus) javanus (Boettger): Ahl, 1931:148. Rhacophorus (Rhacophorus) barbouri: Ahl, 1931:156. Rh. schlegelii margaritifer (Schlegel): Wolf, 1936:187. Rhacophorus javanus (Boettger): Liem, 1971:153. Rhacophorus margaritifer (Schlegel): Frost, Iskandar (1998) examined the type of Rhacophorus depressus and concluded that it is a ranid and not a rhacophorid. Liem (1971) and Wolf (1936) thought that

31 76 HERPETOLOGICAL MONOGRAPHS [No. 16 Rhacophorus moschata Kuhl and van Hasselt was a synonym of R. margaritifer. However, Brongersma (1942) argued that Kuhl and van Hasselt applied this name to a juvenile R. reinwardtii. On 13 May, 1996, Harvey and Smith collected six Rhacophorus margaritifer (UTACV ) at the Taman Safari park near Cibodas, Jawa Barat, Java, Indonesia, S, E. Short descriptions of this species have been published previously (Iskandar, 1998; Liem, 1971; Van Kampen, 1923). Dunn (1928) described the call and color in life of R. margaritifer from Tjibodas (Cibodas, Jawa Barat). Schijfsma (1932) described the tadpole of R. margaritifer. Description and sexual dimorphism. The description is based on five male and one female specimens. Except where noted, morphometric characters of the female fall into the ranges of males, however, means and standard deviations are based on measurements of the males only. With a SVL of mm, our female specimen (Fig. 14) is much larger than the males, which range from mm. This species has a large head % ( ) as wide as long and accounting for % ( ) of SVL. The canthi rostralis and rostral ridges are rounded and low, and the area between the rostral ridges is only weakly depressed. The snout is subacuminate in dorsal view and rounded in lateral view (Fig. 4). In both sexes, an inconspicuous swelling is present at the terminus of the rostral ridges. The lores are weakly concave and the lips are not flared. Eye-nostril distance accounts for % ( ) of the distance from the anterior border of the eye to the tip of the snout, and the nostril is % ( ) closer to the tip of the snout than to the eye. The eye is directed anterolaterally, and the upper eyelid is % ( ) as wide as the interorbital region. A moderate supratympanic fold extending from the posterior edge of the eyelid to the posterior edge of the scapula overlaps the upper edge of the tympanum. The tympanic annulus is visible, though somewhat indistinctly. Length of the tympanum and its FIG. 15. Hand and foot of Rhacophorus margaritifer, an adult female, UTACV annulus is % ( ) as large as eye-diameter. A few low tubercles are present in the folded skin just posterior to the rictus and tympanum. The dorsum, limbs, gular region, and chest are smooth. Posterior to the coracoids, the belly is coarsely areolate, and the ventral surfaces of the thighs are weakly areolate. The supracloacal area bears a crenulated ridge. Numerous, large white tubercles are present below the vent. The arms are short and slender, and the length of the hand is % ( ) of the length of the foot. The fingers are slender and bear truncate (fingers 2 4) or round (finger 1) disks and oval digital pads surrounded by marginal grooves (Fig. 15). Except for slight depressions where the distal phalanges bifurcate, the dorsal surfaces of the digital tips are unmodified. When adpressed, length of finger Webbing between fingers 1 and 2 is evident, though basal. In males variation

32 2002] HERPETOLOGICAL MONOGRAPHS 77 in webbing between the remaining digits is summarized by II( ) (2.5 3 )III(2 2) (2 2)IV. In the female, webbing is slightly more extensive along the fourth finger where 1.5 phalanges are free. Subarticular tubercles are large and round in frontal section, subconical in sagittal section. Round and flat supernumerary tubercles are present proximal to the subarticular tubercles in all specimens. Usually two are present on digits 3 and 4 and one on digits 1 and 2. UTACV has two supernumerary tubercles on each digit. The palmar tubercle is irregular, flat, and bifid. The thenar tubercle is oval and slightly larger than the palmar tubercle. A narrow keel extends along the postaxial edge of finger 4 and the hand. A short space at the wrist separates the keel from a mostly smooth and wider ulnar ridge. No flap or ridge is present at the elbow. The hind limbs are long and slender; TL is % ( ) as long as SVL, and the foot is % ( ) as long as the shank. In males, variation in pedal webbing is summarized by I1 ( )II1 (2-2)III1 (2 2 )IV(1.5 2 ) 1V. The female has slightly more extensive webbing along the preaxial edge of toe 2 (one free phalanx) and toe 3 (1.75 free phalanges). Decreasing lengths of the toes are The disks are subtruncate (toes 2 5) or round (toe 1) and bear oval pads surrounded by marginal grooves. On the foot, subarticular tubercles are large and round in frontal and sagittal section, and proximal subarticular tubercles are about the same size or slightly smaller than distal subarticular tubercles. When present (two of five male specimens), one to three supernumerary tubercles of the foot are flat, round to oval, and indistinct. They are located proximal to the subarticular tubercles on the postaxial digits. The inner metatarsal tubercle is flat, oval, and smaller than the disk of the fourth toe. An outer metatarsal tubercle is absent. A moderate keel extends along the postaxial edge of the fifth toe and is continuous with a weakly crenulated, narrow ridge along the postaxial edge of the tarsus. A subtriangular flap-like calcar is present at the heel. The rictus of UTACV was cut to expose the roof of the mouth. In this specimen, low dentigerous processes of the vomers are located near the anterior edges of the choanae and bear 4/6 subconical teeth. They are about one-third as large as the choanae and are slightly wider than long. At the midline, the processes are separated by a space equal to two times their width. Each process is directed posteromedially at about 10(to the transverse plane. The dentigerous processes and choanae are positioned far laterally. However, the relatively large choanae of this species are clearly and entirely visible at the edges of the lingual shelves of the maxillae when the specimen is on its back. The tongue is subtriangular, deeply notched, and free for about one-third of its length. Paired vocal apertures are elongate slits, about as wide as the middle of the tongue, and visible in the buccal epithelium between the tongue and rictus. In preservative, males have a tan, brown, or gray dorsal ground color. In three of the five males, black specks are more or less uniformly superimposed on the ground color; two other specimens have very few black specks, and those that are present are mostly on the lateral surface of the body. All of the male specimens bear an interorbital pinstripe extending transversely from the lateral margins of the eyelids. Most (four of five) have similar pinstripes between the canthi and on the nape. Irregular, large darker brown blotches mark the dorsa, and 5 6 (counted on the shank) brown bands cross the legs. In males, the dorsal surfaces of fingers 1 2 and toes 1 3 are pale orange, and the remaining dorsal surfaces of the hands and feet are the same color as the dorsum. Webbing is nearly immaculate except between the most postaxial pair of digits on both the hand and foot where the webbing is heavily pigmented. In the female, the dorsal surfaces of the hands and feet and the webbing is gray. UTACV has four small circular white spots on its dorsum. The dorsum of the female is uniformly gray except for about 10 small circular white spots mostly located in the pelvic region but also present on the back,

33 78 HERPETOLOGICAL MONOGRAPHS [No. 16 left forearm, and left thigh. The female lacks pinstripes on its head and lacks brown blotches on its dorsum. Bands on the limbs of the female are indistinct. In both sexes, the supracloacal fold and the ulnar and tarsal ridges are white. The groin is unpatterned and cream; the back of the thigh is immaculate pale orange (most males) or has irregular charcoal blotches (UTACV and the female specimen). A narrow band of white with small charcoal flecks and blotches forming a diffuse net-like pattern separates the brown dorsal pattern from the immaculate cream (gray in the female) venter. The chest and gular regions are immaculate white in males and dirty cream in the female. In life (based on field notes taken by M. B. Harvey for UTACV 54005), the iris is uniformly yellow above and below the pupil. The palpebrum is clear. The legs, arms, hands and feet (including the web) are light orange. The narrow band on the flanks is charcoal and yellow, and the venter is cream. Internal anatomy (based on UTACV 54007, an adult male). The portion of the m. intermandibularis overlapping the m. submentalis is thickened and forms a pair of distinct anterolateral elements (sensu Tyler, 1971b). The median aponeurosis of the m. intermandibularis is round and occupies the middle one-third of the muscle. It does not reach the margin of the m. interhyoideus. The mm. geniohyoidei do not contact one another medially except above the m. submentalis. With the exception of a few diffuse patches of white pigment in the ventral parietal pericardium, the peritonea of this species are not white. The Wolffian ducts are weakly convoluted with indistinct branches extending to the anterolateral portions of the kidneys. A bottle-shaped vesicula seminalis is absent (Bhaduri and Mondal, 1965). In life, the bones were white. The testes are black and much smaller than the trilobed liver. Dermal Roofing Bones: The halves of the sphenethmoid are fused dorsally and ventrally (Fig. 16). The dorsal sphenethmoid is hexagonal, slightly concave, and heavily exostosed. The ventral sphenethmoid is ossified and heavily exostosed; cartilaginous remnants edge the anterior, dorsal, and ventral margins of the optic foramina. The nasals are antero-laterally directed and only slightly overlap the antero-lateral corners of the sphenethmoid. They bear a slender maxillary process extending postero-laterally to overlap the planum antorbitale. The frontoparietals are roughly rectangular in shape. They almost fuse medially: a thin broken fenestra separates their anterior three-fourths. The frontoparietals bear a narrow lateral shelf extending into the orbit to overlap the dorsal edge of the cartilaginous portion of the sphenethmoid. This flange also extends anteriorly to overlap the posterior sphenethmoid. Most of the postero-lateral border of the frontoparietals is fused to the otoccipital. Nasal Capsules: The nares are laterally oriented. The cavum principale is partially enclosed by the tectum nasi, septum nasi, alary cartilage, oblique cartilage, and solum nasi. A long inferior prenasal cartilage extends from the solum nasi to brace against the pars palatina of the premaxilla. Behind the nasal capsules, the vomer is exposed by a large rectangular fenestra and is visible in profile. The superior prenasal cartilage extends from the alary cartilage to brace against the dorso-posterior face of the alary process of the premaxilla. The vertically oriented alary cartilage forms the anterior wall of the narial aperture. The lamina inferior extends dorsally from the crista subnasalis to approach the closed end of the septomaxilla. The crista subnasalis projects anteriorly just above the maxilla and extends ventrally to abut the anterior end of the pars facialis and pars palatina of the maxilla. The capsule bears two short, postero-lateral projections: one extending from the oblique cartilage and one extending from the solum nasi. A narrow planum transversale extends from the posterior edge of the tectum nasi beneath the nasal and connects to the planum antorbitale. The planum antorbitale is partially overlapped by the maxillary process of the nasals and the preorbital process of the maxilla. The planum antorbitale ex-

34 2002] HERPETOLOGICAL MONOGRAPHS 79 FIG. 16. Skull and pectoral girdle of Rhacophorus margaritifer, UTACV 54007, and adult male. Bone is stippled and cartilaginous structures are shaded gray. tends posteriorly along the pars palatina and is continuous with the pterygoid process of the quadrate. Small, triangular cartilages extend into the orbits from the postero-lateral corners of the sphenethmoid and are also continuous with the planum antorbitale. Suspensorium: The squamosal is a triradiate bone, and its ventral ramus is directed anteriorly at 46 to the maxillary arcade. The ventral ramus lies lateral to the quadrate and articulates with the posterodorsal margin of the quadratojugal. The otic ramus is about two-thirds as long as the ventral ramus and directly contacts the otoccipital. The zygomatic ramus is onefourth as long as the ventral ramus and tapers to a point just anterior to the tympanic annulus. The pterygoid is another triradiate element of the suspensorium. Its medial ramus is the shortest arm and lies on the ventral surface of the pseudobasal process with no direct otoccipital contact. The posterior ramus is a flat extension that lies medial to the entirely cartilaginous quadrate. The anterior ramus is the longest and extends to form a broad articulation with the maxilla by means of the quadrate s anterior process. Maxillary Arcade: The premaxillae bear 12/14 teeth. Their lingual shelves are moderately developed and widest medially and laterally. The maxilla bears a rectangular pars facialis and a pointed preorbital process that approximates but does not contact the nasal. Near the ventral edge of the pars facialis and immediately lateral to the ventral and anterior margin of the planum antorbitale, the maxilla bears an L-shaped notch. The maxillae overlap the posterior edges of the premaxillae laterally. The pars palatina of the maxilla is reduced to a narrow shelf, and the maxillae bear 66/67 teeth. The quadratojugal overlaps the maxilla medially. Palate: The vomers are widely separated

35 80 HERPETOLOGICAL MONOGRAPHS [No. 16 and articulate with the postero-lateral edge of the solum nasi. Their dentigerous processes bear 6/8 true teeth and only medially contact the sphenethmoid. The vomers do not contact the maxillae or neopalatines. The neopalatines are slender bones that articulate broadly with the sphenethmoid. They lie medial to the planum antorbitale and approach but do not contact the maxillae. The anterior edge of the cultriform process of the parasphenoid is jagged. This process bears a low, broad median keel and extends forward for about three-fourths of the length of the orbit. Two small, distinct ridges are present at the base of the cultriform process. The alae are postero-laterally directed; they approximate but do not contact the pseudobasal processes. Posteriorly, the parashenoid ends in a rounded point and is not fused to the otoccipital. Mandible: The mentomeckelian bones are ossified and bear a synchondrotic association medially. Laterally, they are fused to the dentaries. The dentary extends posteriorly for approximately half the length of the mandible, covering the dorso-lateral surface of Meckel s cartilage. The angulosplenial spans nearly the entire length of the mandible and covers the ventro-medial surfaces of Meckel s cartilage. It does not contact the dentary or mentomeckelian bone. Meckel s cartilage is entirely cartilaginous except for a small area at the articular surface of the mandible where Meckel s cartilage articulates with the mentomeckelian bone. The angulosplenial bears a broad, flat prominence anterior to the articular surface of the mandible. Hyoid: The hyale are long, slender, and recurved posteriorly; they lack anterior processes. The hyale are slightly swollen at mid-length. The antero-lateral processes are peg-like spikes. The postero-lateral processes are the same shape and approximately twice the length of the antero-lateral processes. The postero-medial processes were damaged during preparation, but appear to directly abut the hyoid plate with no associated cartilaginous stalks. No parahyoid bones are present. Pectoral Girdle: The clavicle lies ventral to an almost completely ossified procoracoid. Coracoids occupy most of the epicoracoid cartilage and are exostosed medially and laterally. The clavicle and coracoid are not fused medially. Proximally, the scapula is bicapitate. The anterior head is fused to the coracoid and clavicle; all three bones are heavily exostosed in this region. The posterior head is fused to the dorso-lateral surface of the coracoid. The suprascapula is fan-shaped and heavily ossified with cartilage being present at its postero-distal borders and a small region posterior to the cleithrum. The antero-distal edge of the suprascapula bears a rounded hooked projection. Relative to the other species, the cleithrum is small and rectangular. It overlaps the anterior border of the suprascapula and terminates short of the scapula-suprascapula articulation. The omosternum is broadly forked posteriorly and bears a large tabular cartilage anteriorly. It articulates with anterior projections of the procoracoid. The sternum is anteriorly dilated and extends posteriorly to expand into a large tabular cartilage. A large fenestra is present in this tabular cartilage, and the base of the tab is ossified. Pelvic Girdle: The iliosacral articulation is type IIB of Emerson (1979). An oval sesamoid bone about one-fourth as long as the sacral diapophysis is present in the transverse ligament. The ilia bear cartilaginous tips at their anterior ends. The ilial shaft is long with a large dorsal crest becoming reduced anteriorly and bearing a rounded prominence dorsal to the acetabulum. The ischium is ventrally fused to the pubis and dorsally fused to the ilium. The pubis is indistinguishable from the ilium and ischium. Postcranial Axial Skeleton: Rhacophorus margaritifer has eight nonimbricate, procoelus, presacral vertebra. The atlantal cotyles are widely spaced. The transverse processes of presacral II are peg-like and bear anterior spikes near their lateral ends. The transverse processes of presacrals II IV, and VIII are perpendicular to the axial column, and transverse processes of the remaining vertebrae are directed posterolaterally. Presacral III and IV bear broad cartilaginous tips that are directed poste-

36 2002] HERPETOLOGICAL MONOGRAPHS 81 FIG. 17. Rhacophorus pardalis (UTACV 54011, SVL 46.31) from 56 km west of Sungaipenuh, Sumatra. riorly, and V VIII are peg-like and bear small cartilaginous tips. The sacral diapophyses are slightly directed posteriorly. The neural arches of presacrals I IV are widely spaced in comparison to the highly overlapping seventh, eighth, and sacral vertebrae. No fusion of elements is present; the neural arches and centra but not the transverse processes are exostosed. Small neural spines are present only on presacrals II and III. The neural arch of the sacral vertebra bears a low horizontal ridge. The coccyx bears a bicondylar association, a dorsal crest on its anterior half, and a large prominence at its anterior terminus. The posterior tip of the coccyx is cartilaginous. No vestigial transverse processes of the coccyx are present. Hand: Rhacophorus margaritifer bears five mesopodial elements in the hand. Two elements are associated with the radioulna. Presumably the postaxial element is the ulnare and the preaxial element is the radiale. The other elements are of uncertain homology. A large cubodal element lies distal to the radiale and ulnare. Two other small elements are proximal to metacarpal I. The prepollex consists of 2 components; both are ossified with the distal constituent being wedge-shaped and bearing a cartilaginous distal edge. Metacarpal III bears a bony prominence along the preaxial margin of its distal end. Metacarpal II bears a smaller but similar prominence in the same location. No sesamoid bones are present in the hand. The terminal phalanges of the hands and feet are Y-shaped. The palmar phalangeal formula is Feet: The feet possess four mesopodial elements. The tibiale and fibulare are fused proximally and distally. An irregularly shaped element of unknown homology lies postaxial to the prehallux and proximal to metatarsal I. A horizontally elongated element of unknown homology lies proximal to metatarsals II and III. The prehallux consists of 2 components, both of which are mostly ossified with cartilaginous distal edges. A small oval sesamoid is present ventral to the proximal end of the tibiale and fibulare. A similar sesamoid is present below the articulation of the femur and tibiafibula. The phalangeal formula of the foot is Natural history. At the Taman Safari park, male Rhacophorus margaritifer called from leaves 1 2 m above ground in the forest near a rocky stream. When prodded with a probe, UTACV lifted up its hands to cover its eyes and slightly flexed its back in an unken reflex. Rhacophorus pardalis Günther Fig. 17 Rhacophorus pardalis Günther, 1858:83. Syntypes BMNH (6 specimens) from Philippines and Borneo. Rhacophorus rizali Boettger, 1897:1. Holotype MTKD 1537 from Dapitan, Mindanao, Philippines. (Note: the holotype is destroyed, Obst, 1977). Rhacophorus pulchellus Werner, 1900:495. Holotype: NHM 1187 from Djapura Sumatra. Polypedates pardalis (Günther): Taylor, 1920:281. Rhacophorus (Rhacophorus) pardalis (Günther): Ahl, 1931:160. Rhacophorus (Rhacophorus) pulchellus (Werner): Ahl, 1931:166. Rh. pardalis pardalis (Günther): Wolf, 1936:204. Rh. pardalis rhyssocephalus Wolf, 1936: 206. Holotype MTKD 786 from Sangi. (Note: the holotype is destroyed, Obst, 1977).

37 82 HERPETOLOGICAL MONOGRAPHS [No. 16 Rh. pardalis pulchellus (Werner): Wolf, 1936:207. Harvey and Smith found an adult male (UTACV 54011, Fig. 17) on vegetation 2 m above ground near a river through forest, 56 km (by road) west of Sungaipenuh on the road to Tapan, Sumatera Barat, Sumatra, Indonesia, S, E, 80 m, C, hrs, 9 June Modern descriptions of this species have been published by several authors (Brown and Alcala 1994, Inger, 1966, Dring, 1979). Here, we focus on characteristics known to be variable among populations and characteristics not mentioned by earlier authors. The specimen is an adult male with a subacuminate and truncate snout. Proximal subarticular tubercles on the third finger are small, oval, and barely discernable; the fourth finger lacks a proximal subarticular tubercle. Webbing is typical for the species; web reaches the disks of all digits except along the preaxial edge of finger 2 and postaxial edge of finger 1 where it extends slightly beyond the subarticular tubercle. On both hands, a round and low metacarpal supernumerary tubercle is present on finger 2. The skin is coarsely granular on the ventral body, weakly so on the ventral surface of the thigh and gular region, and smooth on the chest. The nuptial pads are an elongate strip along the preaxial edge of the first finger from just distal to the subarticular tubercle to about the middle of the thenar tubercle. Nuptial excrescences cover only the edge of the metacarpal segment on the dorsal side. The specimen has faded little since preservation. The dorsum is grayish tan with black speckling. A circular cream spot, edged in black, and a little larger than the tympanum is present above the distal end of the urostyle; a larger cream spot with black edging overlaps the distal shank and proximal tarsus. Banding on the limbs is very faint; two bands cross the forearm and three cross the shank. In life, the pale yellow of the venter and gular regions graded to bright yellow on the flanks; both areas are now white. Ventral surfaces of the thighs and arms are peach. Black reticulation covers most of the belly and extends high onto the flanks. Black reticulation also extends along the ventro-postaxial edge of the antebrachium, hand, shank, and foot. Eight small black spots lie below the mandible. The sides of the hind limbs are immaculate orange. In life, webbing of the hands and feet was red-orange. The iris was orange, and the ciliary ring narrow and yellow. The palpebrum was clear with an indistinct distal edge and the conjunctiva was gray. The Wolffian ducts are convoluted with short processes extending to the kidneys. A bottle-shaped vesicula seminalis is absent (Bhaduri and Mondal, 1965). A few white fibers are present in the parietal peritoneum below the m. rectus abdominis, otherwise the peritonea are not white. The testes are cream and much smaller than the trilobed liver. Measurements (in mm) and morphometric ratios: SVL 46.31, HL 14.8, HW 15.52, ED 6.44, EN 4.86, NS 2.74, TympL 2.88, IOD 4.52, EW 5.14, IND 3.63, HL 13.38, FL 22.73, TL 25.04, DF3W 2.72, DF1W 1.55, DT4W HL/SVL 0.32, HW/HL 1.05, NS/EN 0.57, EW/IOD 1.14, IND/ IOD 0.80, TympL/ED 0.45, HL/FL 0.59, FL/TL 1.10, DF1W/DF3W 0.57, DT4W/ DF3W 0.68, DF3W/TympL Remarks. We agree with the opinion of earlier authors (Inger, 1954, 1966; Wolf, 1936) that Rhacophorus pulchellus Werner is a synonym of R. pardalis. The holotype (NHM 1187) has faded to dark brown and pattern can barely be discerned. More metacarpal supernumerary tubercles are present in Werner s specimen than in ours (UTACV 54011), 1 3 being present on each of the fingers. Although this species usually lacks a proximal subarticular tubercle on the fourth finger, the type has a very small, oval one on both hands. Three bands cross the forearm and four cross the shank of the type. Rhacophorus prominanus Smith Fig. 18 Rhacophorus prominanus Smith, 1924: 185. Holotype BMNH (formerly M.

38 2002] HERPETOLOGICAL MONOGRAPHS 83 FIG. 18. Rhacophorus prominanus (UTACV 54012, SVL mm), an adult male from Gunung Rajabasa, Sumatra. Smith 6691) from Jor, Batang Padang, Perak, altitude 600 meters, Malaysia. Rhacophorus (Rhacophorus) prominanus (Smith): Ahl, 1931:165. Rhacophorus dulitensis [in part]. Wolf, 1936:210. Rhacophorus dulitensis prominanus Taylor, 1962:491. Rhacophorus prominanus (Smith): Inger, 1966:295. Rhacophorus tunkui Kiew, 1987:418. Holotype UMKL 1986 from Sungai Jasin, Ulu Endau, Johore, Malaysia. New synonymy. Justification of synonymy. Kiew (1987) assigned a new name to a frog collected at Sungai Jasin, Ulu Endau, Johore (his holotype, UMKL 1986) and specimens (BMNH ) identified as Rhacophorus prominanus by Berry (1975) and collected at Kuala Tahan, Taman Negara, Pahang, Malaysia. Kiew writes that his putative new species differs from Rh. prominanus in being smaller in size with a shorter and more lightly built body (Kiew, 1987:421). A lightly built body is a consequence of smaller size. A photograph (his plate 1) comparing the holotype and a specimen of Rhacophorus prominanus appears to illustrate this difference. However, like most of its congeners, females of R. prominanus are considerably larger than males. Kiew fails to report the sex of any of his type series, including the holotype, nor does he explain how he determined his holotype was an adult. No mention of nuptial excrescences appears in his description. The specimen on the left of Kiew s plate 1 is comparable in size to females of Malaysian R. prominanus examined by us (e.g., FMNH with a SVL of 47.61), and this plate appears to contrast a male and female specimen of the same species. Without exception, snout shape was sexually dimorphic among Rhacophorus examined by us. Failure to consider sexual differences may also have led Kiew to conclude that the snout of his putative species has a more acute slope at the canthus and is shorter than that of Rh. prominanus (p. 421). Kiew writes that the back of his putative new species is less densely spotted with pigment (p. 421) than Rhacophorus prominanus. Spotting in both R. prominanus and R. dulitensis is more variable than the author realizes. Among the specimens of R. prominanus examined by us, FMNH has almost no dorsal spots, FMNH is heavily spotted though not as much as the female in Kiew s plate, and the remaining specimens have relatively few spots. Sumatran specimens (Fig. 19) exhibit similar extremes in degree of spotting. Kiew writes that his putative new species has a more distinct canthus rostralis demarcated by pigmentation (p. 421). His meaning is clarified in the diagnosis, canthus rostralis distinct and demarcated by a white pigmented line accompanied ventrally by a brown line (p. 419). However, these same words describe the canthus of most Rhacophorus prominanus and R. dulitensis. In these species, a sharp white line usually extends from the tip of the snout, above the nostril and just above the canthus to the orbit (Fig. 18, see also figure 104 of Inger and Stuebing, 1997). The white line is bordered ventrally by dense melanophores. Although usually present, this line may be incomplete and indistinct or absent in occasional specimens (e.g., FMNH ). Finally, Kiew writes, the skin flaps over the anus, at the heels, and along the outer

39 84 HERPETOLOGICAL MONOGRAPHS [No. 16 FIG. 20. Habitat of Rhacophorus prominanus, a mountain stream near Kalianda, southern slope of Gunung Rajabasa, Sumatra, 420 m. FIG. 19. Rhacophorus prominanus from Sumatra (UTACV 54012, top specimen, SVL mm, and UTACV 54013). margin of the lower arm and metatarsus are weaker (p. 421) in his putative new species. Neither his description nor the poor illustration of the holotype (his figure 1) clarifies what Kiew might have meant by weaker. Nonetheless, width of these folds varies considerably within species and all of them tend to be thicker and wider in larger specimens. In summary, putative differences between Kiew s species and Rhacophorus prominanus are the result of sexual dimorphism and individual variation within a single species. Accordingly we consider Rhacophorus tunkui Kiew to be a junior synonym of Rhacophorus prominanus Smith. Harvey and Smith collected two adult male (Fig. 19) Rhacophorus prominanus. One specimen (UTACV 54012) was in vegetation above a shallow pool near a river (Fig. 20) between hrs and air temperature of 21 C on the southern slope of Gunung Rajabasa, Lampung, Sumatra, Indonesia, S, E, 430 m. A second specimen (UTACV 54013) was collected at 700 m along a stream through forest at 2100 hrs and 21 C, 34 km (by road) west of Sungaipenuh on the road to Tapan, Jambi, Sumatra. The specimens confirm Inger s (1966) suspicion that this species would eventually be found in Sumatra. Short and incomplete descriptions of this species have been published for populations in Malaysia (Berry, 1975) and Thailand (Taylor, 1962). Chan- Ard et al. (1999) provided color photographs of male and female Malaysian specimens. Description. Both specimens are adult males. Their heads are % as wide as long and account for 27 28% of SVL. The snout is acuminate in dorsal view, spatulate and sloping in profile. Low rostral ridges enclose a depressed area in front of the nostrils. The nares are 75 87% closer to the eye than to the tip of the snout, and IND is 73 79% of IOD. The canthus rostralis is rounded and straight; the lores slope to the mouth, and the lips are not flared. The eyes are directed anterolaterally; the upper eyelid is 73 79% as wide as the interorbital space. A very low and inconspicuous (most easily seen in UTACV 54012) supratympanic fold ex-