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On Leptobrachium from Thailand with a Description of a New Species (Anura: Pelobatidae)

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On Leptobrachium from Thailand with a Description of a New Species (Anura: Pelobatidae) Japanese Journal of Herpetology 18 (1): 19-29., June 1999 (C)1999 by The HerpetologicalSociety of Japan On Leptobrachium from Thailand with a Description of a New Species (Anura: Pelobatidae) MASAFUMI MATSUI, JARUJIN NABHITABHATA, AND SOMSAK PANHA Abstract: We compared morphological characters among Thai populations of Lepto- brachium currently assigned to L. hasseltii or L. pullum. We also examined syn- types of the latter, which was first described from Vietnam as Megalophrys hasseltii var. pullus Smith, 1921, and designated the lectotype. The Thai populations are slightly variable in adult and larval morphology, but are collectively different from L. hasseltii (originally described from Java) or L. pullum. We therefore describe a new species, L. smithi, for those populations on the basis of the sample from Khao Chong, southern Thailand. Key words: Pelobatidae; Leptobrachium; Systematics; Thailand; Biogeography The genus Leptobrachium (sensu stricto [not 239) and Manthey and Grossmann (1997) clearly of Dubois, 1980]=subgenus Leptobrachium of presented through photographs of individuals Dubois and Ohler, 1998) is a group of pelobatid from western and northeastern Thailand, frogs characterized by a stocky body with slen- respectively. der, short hindlimbs (Inger and Stuebing, 1997; However, Smith (1921) also clearly indicated more detailed diagnosis in Dubois and Ohler, that he split M. h. var. pullus from "typical has- 1998). It occurs from southern China to the seltii" from Siam (=Thailand) and the Malay Philippines, Indonesia and the Sunda Is. to Bali Peninsula on the basis of the differences in body (Frost, 1985). The type species of this genus, L. shape and coloration. Because he is famous for hasseltii, has long been regarded as widely dis- his very active surveys throughout Thailand tributed throughout the range of the genus (e.g., (e. g., Smith, 1922) and must have been very Taylor, 1962; Inger, 1966), although most of the familiar with Leptobrachium from this country, previous authors surmised this apparently heter- the above statement by him deserves special at- ogeneous species to be a composite of several tention. The current taxonomic arrangement of forms. Taylor (1962), for example, suggested applying the name of L. pullum to the Thai occurrences of distinct forms within Thailand. populations simply on the basis of similar eye Few authors, however, seriously studied these color is logically problematic. Nevertheless, taxonomically problematic frogs until recently very few recent authors have actually compared (detailed review in Dubois and Ohler, 1998). the Thai populations of Leptobrachium with the Inger (1983), following personal information typical L. pullum in detail (e. g., Inger et al., from Julian Dring, adopted the name L. pullum 1999). for the population from southern Thailand, and During our five years' field survey of many Dubois and Ohler (1998) also tentatively referred places in Thailand, we observed a number of to populations from Thailand, as well as from adult and larval specimens of Leptobrachium in Myammar and Vietnam by this name. Lepto- life and recorded their calls. One of us (MM) brachium pullum was originally described by also examined syntypes of Megalophrys hasseltii Smith (1921) as Megalophrys hasseltii var. pullus var. pullus Smith, 1921 (= Leptobrachium pul- from Arbre Broye, Langbian Plateau, Southern lum) stored at the Natural History Museum, Annam (Vietnam). Smith (1921:423) reported U. K. (BM) and of Leptobrachium hasseltii kept that the upper half of the iris of this species in at the Museum of Natural History at Leiden life is scarlet. Recent recognition of the Thai (RMNH). populations as L. pullum probably relies on this In this paper, we analyze variations among the iris color, which Zhao and Adler (1993, as L. Thai populations and also compare them mor- hasseltii: commented by Matsui and Ota, 1995: phologically with syntypes of L. pullum and L. hasseltii. As a result, we describe a new species Accepted 13 May 1999 from Khao Chong, southern Thailand. 20 Jpn. J. Herpetol. 18 (1) 1999 Nakhon Si Thammarat Province, alt. 370m; MATERIALS AND METHODS (11) Khao Pra Taw, Phuket, Phuket Province, Extensive field surveys were made throughout alt. 80m; (12) Khao Chong, Trang Province (at Thailand between September of 1993 and Sep- Ton Nam Plu Waterfall, alt. 260m, Ton Te tember of 1997 (Fig. 1). We collected speci- Waterfall, alt. 210m, and Sairung Waterfall, mens, fixed them in 10% formalin, and later alt. 260m). We could not find Leptobrachium stored them in 70% ethanol. Whenever possi- in other parts of Thailand, notably in the eastern ble, we recorded calls using a cassette tape and central regions (Fig. 1). recorder (Sony TC-D5) with an external micro- Adult morphology. -In all specimens ob- phone (Sony ECM-23), and later analyzed them served except one from Ban Khun Klang, Doi using computer programs, SoundEdit Vers. 2 Inthanon, the upper half of the iris was scarlet, and SoundEdit Pro (MacroMind-Paracomp, orange, or yellow. The exceptional individual Inc.) on a Macintosh computer. Terminology with the upper half of the iris light sky blue was for acoustic characteristics follows Matsui (1997). For all preserved adult specimens, we took the following eight body measurements following Matsui (1984): 1) snout-vent length (SVL); 2) head length (HL); 3) head width (HW); 4) lower arm length (LAL); 5) tibia length (TL); 6) foot length (FL); 7) hindlimb length (HLL); 8) inner metatarsal tubercle length (IMTL). All meas- urements were made to the nearest 0.1mm with a dial caliper under a binocular dissecting micro- scope. Larvae were assigned to developmental stages following Gosner (1960) and measured following Matsui (1979). Variation in SVL was examined by analysis- of-variance (ANOVA) with the Tukey range test. Each of the other characters was converted to a percentage ratio (R) in relation to SVL and statistical comparisons were made among popu- lations with a sample size of four or larger when males and females were combined. For these variables, Kruskal-Wallis tests with nonparamet- ric multiple comparisons or Mann-Whitney U tests were performed to detect the presence or absence of differences in the frequency distribu- tions. The significance level was set at 0.05. RESULTS Distribution of Leptobrachium in Thailand. - From our field surveys, we confirmed the occur- rences of Leptobrachium at the following locali- ties: (1) Phu Luang, Loei Province, alt. 980m; (2) Mae Hong Son, Mae Hong Son Province (at Pha Sua Waterfall, alt. 570m); (3) Doi Suthep, Chiang Mai Province (at Montathan Waterfall, FIG. 1. Map of Thailand, showing sampling local- ities where specimens of Leptobrachium were sampled alt. 1000m); (4) Doi Inthanon, Chiang Mai (closed circles) or not sampled (open circles). A: Phu Province (at Ban Khun Klang, alt. 1350m); (5) Luang, Loei; B: Mae Hong Son, Mae Hong Son; C: Sangkhla Buri Kanchanaburi Province, alt. Doi Sthep, Chiang Mai; D: Doi Inthanon, Chiang 540m; (6) Erawan, Kanchanaburi Province, alt. Mai; E: Sangkhla Buri, Kanchanaburi; F: Erawan, 240m; (7) Pilok, Kanchanaburi Province, alt. Kanchanaburi; G: Pilok, Kanchanaburi; H: Pa Lao U, 620m; (8) Pa Lao U, Phetchaburi Province, alt. Phetchaburi; I: Khlong Saen, Surat Thani; J: Khao 280m; (9) Khlong Saen Wildlife Sanctuary, Sur- Luang, Nakhon Si Thammarat; K: Phuket, Phuket; L: at Thani Province, alt. 130m; (10) Khao Luang, Khao Chong, Trang. MATSUI ET AL. -NEW PELOBATID FROG 21 identified as L. chapaense (see Dubois and Oh- ler, 1998), and was excluded from the analyses. Of the remainder, the population from Khao Luang, Nakhon Si Thammarat had an iris with a yellow upper half, whereas the irises of the other populations were scarlet or orange dorsally. Of the two specimens collected at exactly the same locality in Pa Lao U, Phetchaburi Province, one had yellow and the other orange coloration in the upper half of the iris. A summary of measurements is shown in Ta- bles 1 and 2. No characters examined showed a definite geographic pattern of variation. The population sample of Erawan was the largest in male body size with a mean (±SD) SVL of 60.0±5.7mm (Table 1), and that from Khao Chong was the smallest (mean±SD=41.4 ±2.5mm). The Erawan sample was sig- nifficantly larger than all other samples in SVL (p<0.05), but the Khao Chong sample did not differ from several other samples. Of the remaining characters examined (Table 2), sig- nificant differences (p<0.05) among population samples were found only in RLAL and RHL. Samples from Nakon Si Thammarat and Khao Chong had significantly larger RLAL (medi- ans=56.9% and 56.0% SVL, respectively) than samples from Erawan (50.8% SVL) and Mae Hong Son (51.8% SVL), and the sample from Nakon Si Thammarat had larger RHLL (127.7% SVL) than that from Mae Hong Son (118.0% SVL). No other combinations or characters yielded significant differences. Body coloration was also variable, with the TABLE1. Morphometric variation in SVL (in mm) in Leptobrachium from Thailand and L. pullum. 22 Jpn. J. Herpetol. 18 (1) 1999 ground color of the dorsum varying from gray to them in designating the lectotype other than dark purplish brown. Most specimens from choosing a specimen from the type locality Erawan had a white ventrum with dark markings designated by Smith (1921:440). We hereby only on the chin and around the groin, whereas designate a male specimen BM 1921. 5. 5. 36 (No. the ventral sides of specimens from other locali- 5192 of Smith, 1921:440) from Arbre Broye as ties were white with varying amounts of darker lectotype of Megalophrys hasselti var. pullus infusion. Smith, 1921 (Fig. 5). Tadpoles. -We were able to find tadpoles of Smith (1921) noted that M. h. var. pullus Leptobrachium only in the southern localities (Pilok, Khao Chong, and Phuket). Larvae from these three localities were similar in color pattern to that described by Inger (1983) for the larvae tentatively assigned to L.
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