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Home , Buergeria, Chirixalus, Chiromantis, Feihyla, Ghatixalus, Kurixalus

Asian Herpetological Research 2011, 2(1): 1-11 DOI: 10.3724/SP.J.1245.2011.00001

The Systematic Status of Rhacophorid

Jiatang LI1, 2, Dingqi RAO2, Robert W. MURPHY2, 3 and Yaping ZHANG2*

1 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China 2 State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, Yunnan, China 3 Centre for and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada

Abstract The family recently underwent notable reorganization and taxonomic rearrangement. Several new genera were erected to reflect evolutionary history, including , Gracixalus, , and . Herein, we reviewed the systematics and phylogeny of the rhacophorids, the definition of the family Rhacophoridae and its tribes, and the taxonomic history and diagnoses of the genera of this family. Also, we suggest future directions for research. Keywords progress, prospect, faunal survey, systematics, , China

1. Introduction from living on soil to the forest canopy (Amphibian Web, 2010). Some interesting reproductive modes occur Today, the family Rhacophoridae consists of 319 in rhacophorids. Several genera, including , in 14 genera (Biju et al., 2010; Frost, 2010). The group and Raorchestes, are characterized by ranges across tropical Africa and Southern Asia, including the aerial direct development of eggs into froglets (Bahir India and Sri Lanka, the Himalayas, , South et al., 2005; Biju et al., 2010; Bossuyt and Dubois, 2001; China, eastward to Japan and the Philippine Islands, Dring, 1979; Dutta, 1997). The genera , and southward across Indochina to Greater Sunda Feihyla, and either form (Figure 1) (Frost, 2010; Liem, 1970). Most rhacophorids aerial nests of foam or they lay their eggs in a jelly-like are treefrogs, with the majority of species exhibiting secretion filled with bubbles which hangs over water. particularly vivid and pleasant coloration. The species are Their eggs develop into tadpoles that drop into the water united morphologically in possessing intercalary elements once they reach a certain size (Bain and Nguyen, 2004; between the two distal phalanges and expanded digit Boulenger, 1903; Duellman and Trueb, 1986; Inger, 1966; discs. These characters are shared with hylids, mantellids, Liem, 1970; Orlov et al., 2002, 2004). Treefrogs are and some ranid lineages (Frost et al., 2006; Liem, 1970; known for their loud and diverse vocalizations, made only Manzano et al., 2007). Toe pads help in climbing due by the males, with flap consonant. to their ectodermal morphology and secreted proteins Rhacophorid has been a matter of much which aid in their adherence to surfaces (Duellman and recent debate. Previous classifications are based on Trueb, 1986). Many treefrogs can change color, usually to either a single morphological character or a complex match their surroundings. When the reproductive season of characters (Channing, 1989; Jiang et al., 1987; has passed, treefrogs usually burrow into the humus on Liem, 1970; Zhao and Adler, 1993). Historically, the forest floor. Rhacophorids range from being small morphological studies played a central role in to large, with SVL (snout-vent length, from tip of snout the development of rhacophorid systematics and to vent) ranging from 15 to 110 mm. The habitats vary taxonomy. Within this body of research, representative studies include Channing (1989), Jiang et al. (1987), * Corresponding author: Prof. Yaping ZHANG, from Kunming Institute Liem (1970) and Wilkinson and Drewes (2000). of Zoology, Chinese Academy of Sciences, with his research mainly Nowadays, DNA sequences are commonly used for focusing on conservation genetics, molecular evolution, and population inferring phylogenetic relationships (Frost et al., genetics. Email: [email protected] 2006; Grosjean et al., 2008; Li et al., 2008, 2009; Received: July 23 2010 Accepted: 30 December 2010 Meegaskumbura et al., 2002; Richards and Moore, 2 Asian Herpetological Research Vol. 2

Figure 1 Geographic distribution of the extant Rhacophoridae shown by the red patches. The Figure is modified from Vitt and Caldwell (2009).

1998; Wilkinson et al., 2002; Yu et al., 2008, 2009) Channing (1989) reanalyzed Liem’s data and and making taxonomic revisions. Stemming from the obtained a different cladogram. His results suggested recent changes, herein we review the phylogenetic that the subfamily Mantellinae, including Mantella and systematics of rhacophorids, the definition of the Laurentomantis, should be moved from the Ranidae to the family Rhacophoridae and its tribes, and the taxonomic Rhacophoridae. history of the genera recognized within this family. We Other studies have focused on rhacophorid systematics. also provide suggestions for future research. Jiang et al. (1987) constructed a phylogenetic hypothesis for Chinese species using morphological characters. 2. Systematics and Phylogeny of Rhacophorids They resolved Polypedates and Rhacophorus as sister taxa, and together these genera formed the sister clade of Ahl (1931), in the first major taxonomic revision of “”. Philautus rooted as the sister group to this the family Rhacophoridae, recognized 12 genera and clade and being the most basal . Using 527 species distributed in Africa, Madagascar, the allozyme data and the incorporation of Japanese species, Seychelles Islands, and Asia. Based on an extensive suite Nishioka et al. (1987) also resolved Buergeria as the most of osteological characters, Laurent (1943, 1946) erected basal genus and Polypedates and Rhacophorus as sister a new subfamily, the Hyperoliinae, to include all of the lineages. Yang and Lin (1997), and Wilkinson (1997) African treefrog genera, except for Chiromantis. The came to similar conclusions in that Rhacophorus and subfamily was considered as the familial rank by Laurent Polypedates were sister taxa followed downward on the (1951) without diagnosed characters. The placement tree by “Chirixalus” and then Buergeria. of Afro-Asian species into the Rhacophoridae, and Wilkinson and Drewes (2000) produced a comprehensive that of those from Madagascar into another family, the phylogenetic reconstruction for the Rhacophoridae based on Mantellidae, have been widely accepted recently (e.g., the data largely taken from Liem’s (1970) and Channing’s Frost et al., 2006; Van der Meijden et al., 2005; Vences (1989) studies. They identified synapomorphies for the and Glaw, 2001). major clades and all genera, and summarized intergeneric Liem (1970) made the first phylogeny-based revision relationships. Their results resolved Rhacophorus as the of the rhacophorids. His study used myological, skeletal, sister group of Polypedates as opposed to “Chirixalus”. and external morphology of 420 species of Old World and Philautus were resolved as sister taxa for treefrogs. Liem defined the major evolutionary lines the first time. All Asian genera, except Buergeria, formed within the assemblage and made some taxonomic a monophyletic group, and Polypedates and Rhacophorus suggestions with respect to genera. He proposed that were sister genera. the Rhacophoridae probably evolved from an Asian Phylogenetic hypotheses continued to differ. ranid stock. One genus appeared to have reached Africa Using 1081 bp of nucleotide sequence data from the relatively recently. This conclusion was supported by Li et mtDNA 12S and 16S genes, Richards and Moore al. (2008, 2009) and Wilkinson et al. (2002). (1998) placed Chiromantis as the sister taxon to No. 1 Yaping ZAHANG et al. The Systematic Status of Rhacophorid Frogs 3

Polypedates. Subsequently, Emerson et al. (2000) Using 1676 bp of mitochondrial and nuclear gene placed Chiromantis as the sister taxon to “Chirixalus”, fragments (12S, 16S and rhodopsin) from 48 rhacophorids, and Richards et al. (2000) resolved it as the sister including 16 undescribed Sri Lankan and southern Indian taxon to Rhacophorus. Also using DNA sequence species, Grosjean et al. (2008) recognized both subfamilies data, Wilkinson et al. (2002) provided evidence for Buergeriinae and and erected the new tribe a sister group relationship of the Rhacophoridae and Nyctixalini. The basal position of Buergeria within the Mantellidae. They considered that these families Rhacophorinae was well supported, and Nyctixalus and should be subfamilies within Rhacophoridae. were sister taxa. The type species of Feihyla Consistent with the previous studies (Channing, (Chirixalus palpebralis) nested at the base of the tree for 1989; Jiang et al., 1987; Liem, 1970; Wilkinson and Chiromantis, albeit, with low support. These authors did Drewes, 2000), Buergeria rooted at the base of the not believe Feihyla deserved recognition; they recognized tree for the Rhacophorinae. Curiously, ‘‘Chirixalus’’ “Chirixalus palpebralis” as belonging to Chiromantis. palbebralis was isolated phylogenetically from their The following sets of relationships were congruent with representative rhacophorids. Further, “Chirixalus” was those of Frost et al. (2006): (1) Chirixalus was a junior not monophyletic, Chiromantis and Polypedates had a synonym of Chiromantis; (2) was closely close relationship, and Rhacophorus formed the sister related to Aquixalus; and (3) Kurixalus idiootocus and K. clade to Philautus plus “Chirixalus”. Nyctixalus and verrucosus were in the correct genus. Adding more data Theloderma were resolved as sister taxa. from India, Sri Lanka and the Sunda Islands, Philautus Delorme et al. (2005) constructed a phylogeny, and was not a monophyletic group. The clades of Philautus proposed a new taxonomy based on morphological and did not coincide with geographic distributions. Grosjean molecular data. However, they did not detail all the et al. (2008) also studied the evolution of reproduction in evidences supporting their phylogeny. Their phylogeny the Rhacophoridae and found that the ability to produce was similar to that of Channing (1989), Liem (1970), a foam nest emerged only once, as previously proposed Jiang et al. (1987), Wilkinson and Drewes (2000), and (Wilkinson et al., 2002). Wilkinson et al. (2002) as follows: Buergeria was the Li et al. (2008) employed maximum parsimony, sister taxon of all the remaining rhacophorids; Theloderma maximum likelihood, and Bayesian inference analyses and Nyctixalus were sister taxa; and Chirixalus was to investigate the molecular phylogenetic relationships not resolved as being monophyletic because of C. of the Rhacophoridae. They used 2904 bp of sequences palpebralis. Surprisingly, Delorme et al. (2005) erected from both mitochondrial (12S rRNA, 16S rRNA, tRNA a non-monophyletic new genus Aquixalus with type for valine) and nuclear (tyrosinase, rhodopsin) genes. species “Philautus” odontotarsus and divided Aquixalus Consistent with the studies by Delorme et al. (2005) and into the subgenera Aquixalus (type species “Philautus” Frost et al. (2006), Aquixalus was not a monophyletic odontotarsus) and Gracixalus (type species “Philautus” assemblage of species. Aquixalus (Aquixalus) formed a gracilipes). Unfortunately, their morphological diagnosis well-supported monophyletic group within Kurixalus, of Aquixalus also applied to some species of Kurixalus. and Aquixalus (Gracixalus) was more closely related They assigned some species of “Philautus” to Aquixalus to the species of Rhacophorus, Polypedates and without reference to a phylogeny. Subsequently, Fei et al. Chiromantis than to Aquixalus (Aquixalus). Consequently, (2009, 2010 a) assigned all Chinese species of “Philautus” they synonymized Aquixalus (Aquixalus) into Kurixalus, to Aquixalus. and elevated the subgenus Gracixalus to full generic rank. Frost et al. (2006) supported Channing’s (1989) After moving R. hainanus to Kurixalus, the monophyly of recognition of the subfamilies Buergeriinae and Rhacophorus was strongly supported. Philautus did not Rhacophorinae. In order to avoid the polyphyly of form a monophyletic group because Philautus (Kirtixalus) the genus Chirixalus, they erected the new genus was the sister group to the clade comprised of Kurixalus Feihyla to contain “Chirixalus’’ palpebralis. Thay also and Aquixalus (Aquixalus). However, Li et al. (2008) placed Chirixalus into synonymy with Chiromantis did not make taxonomic changes in Philautus because and suggested that Kurixalus and Aquixalus were of inadequate taxonomic sampling. Chiromantis romeri morphologically plesiomorphic. They recognized the did not cluster with the other species of Chiromantis, but putatively monophyletic subgenus Gracixalus whose type rather formed the sister group to all other rhacophorids species, G. gracilipes, rooted at the base of their tree for save Buergeria. Because of the phylogenetic position the subfamily Rhacophorinae. of Chiromantis romeri, Li et al. (2008) erected the 4 Asian Herpetological Research Vol. 2 new genus Liuixalus to contain it. They suggested that some bubbles. Terrestrial direct development evolved mainland Chinese populations of P. megacephalus are twice in the Rhacophoridae. likely a complex of at least two species, according to their phylogeny and the studies of Narins et al. (1998), Orlov et 3. Definition of the Family Rhacophoridae and Its al. (2002), and Trépanier et al. (1999). Tribes Yu et al. (2008) used 23 Chinese rhacophorid species in their phylogenetic analyses of mtDNA sequences. Their The name Rhacophoridae Hoffman, 1932 takes the results were largely consistent with those of Frost et al. date of publication for Polypedatidae Günther, 1858, (2006), Li et al. (2008), and Yu et al. (2007). For example, and thus has priority under the provisions of Article 40 “Philautus” romeri was the sister taxon to all other of the International Code of Zoological Nomenclature rhacophorines, and “Chirixalus” doriae nested within (1999). The type genus is Rhacophorus Kuhl et Van Chiromantis. In order to obtain a monophyletic Philautus, Hasselt, 1822. Several studies consider rhacophorids they transferred P. rhododiscus and P. albopunctatus into to be a monophyletic subfamily, the Rhacophorinae, Theloderma, which was also published by Yu et al. (2007). within the Ranidae (Dubois, 1981, 1987; Liem, 1970; Adding several nuclear protein-coding genes, Yu et al. Wilkinson and Drewes, 2000; Wilkinson et al., 2002). (2009) expanded on their hypothesis for the phylogenetic Channing (1989) reanalyzed Liem’s data and divided relationships within the Rhacophoridae. They resolved the Rhacophoridae into two subfamilies, Buergeriinae Chiromantis as the sister group to the clade formed by and Rhacophorinae. Dubois (1992) proposed a major Feihyla, Polypedates and Rhacophorus, and confirmed taxonomic revision, while considering the group to be that Feihyla was a valid genus. They also suggested that a subfamily of the Ranidae. Marmayou et al. (2000) the Chinese Rhacophorus needed revision. Rhacophorus suggested that rhacophorids were deeply imbedded pingbianensis was considered to be a junior synonym within the Ranidae. In their anatomical and molecu- of R. omeimontis. In order to maintain a monophyletic lar analyses, Frost et al. (2006) recognized the family Philautus, Yu et al. (2009) elevated subgenus Kirtixalus to Rhacophoridae and this usage was consistent with most the rank of genus and placed P. menglaensis within it as K. other studies (Bossuyt and Roelants, 2009; Fei et al., menglaensis. 2009; Grosjean et al., 2008; Jiang et al., 1987; Li et Most recently, Li et al. (2009) investigated the al., 2008, 2009; Yu et al., 2008, 2009; Zhao and Adler, phylogenetic relationships among 12 genera of 1993; Zhao et al., 2000). Channing’s recognition of the treefrogs based on a large DNA sequence data set, two subfamilies has been strongly supported by several including five nuDNA (brain-derived neurotrophic molecular studies (Bossuyt et al., 2006; Frost et al., factor, proopiomelanocortin, recombination activating 2006; Li et al., 2008, 2009; Wilkinson et al., 2002; Yu gene 1, tyrosinase, and rhodopsin) and three mtDNA et al., 2009). (partial 12S and 16S rRNA, and tRNA valine) genes. Dubois (1992) recognized three rhacophorid tribes: Analysis of the nuDNA data resolved three major Buergerini, Philautini, and Rhacophorini. Within clades. Gracixalus, Philautus, Theloderma moloch, these tribes, the Buergerini included only one genus, Pseudophilautus, and Kurixalus constituted the Buergeria. The Philautini contained “Aquixalus”, first group with the latter two being sister taxa. The Kurixalus, Nyctixalus, Philautus, and Theloderma, and second group consisted of Nyctixalus and Theloderma the Rhacophorini included Chirixalus, Chiromantis, exclusive of T. moloch. The third group contained Polypedates, and Rhacophorus (Dubois, 2005). Using Feihyla, Polypedates, Rhacophorus and Chiromantis mitochondrial DNA (mtDNA) and nuclear DNA (nuDNA) vittatus. The elevation of Kirtixalus to generic rank sequence data, Grosjean et al. (2008) suggested that (Yu et al., 2009) was justified, but the name Kirtixalus the Rhacophorinae should consist of two tribes, the Dubois, 1987 was replaced with Pseudophilautus Rhacophorini and Nyctixalini, the latter containing Laurent, 1943 because of priority. In order to maintain Theloderma and Nyctixalus. Using an even larger the monophyly of Philautus, Li et al. (2009) transferred DNA sequence dataset, Li et al. (2009) erected the P. ocellatus and P. hainanus into Liuixalus, P. rhacophorine tribe Liuixalini, (type genus Liuixalus) and longchuanensis and P. gryllus into Pseudophilautus, P. recognized the Nyctixalini (Theloderma and Nyctixalus) banaensis into Kurixalus, and P. quyeti into Gracixalus. and Rhacophorini (Chiromantis, Feihyla, Ghatixalus, Again, foam nesting was resolved to have evolved only Gracixalus, Kurixalus,Rhacophorus, Philautus, once, as did lay eggs in a jelly-like matrix containing Polypedates, and Pseudophilautus). The subfamily No. 1 Yaping ZAHANG et al. The Systematic Status of Rhacophorid Frogs 5

Buergeriinae has been maintained without modification northern Viet Nam (Frost, 2010). This reproductive mode, (Channing, 1989; Delorme et al., 2005; Frost et al., 2006; laying eggs in a jelly containing some bubbles, serves as Grosjean et al., 2008; Li et al., 2008, 2009; Liem, 1970; a behavioral synapomorphy to diagnose Feihyla (Li et al., Wilkinson et al., 2002; Yu et al., 2009). 2009). It presently contains F. palpebalis, F. fuhua, and F. vittatus (Fei et al., 2010 b; Frost, 2010). 4. Taxonomic History and Diagnoses of the Ghatixalus was described by Biju et al. (2008) Genera of Rhacophoridae using Polypedates variabilis as the type species. Biju et al. (2008) considered the genus to be the sister The subfamily Buergeriinae contains a single genus, taxon of Polypedates. They assigned two species to the Buergeria, which has Hyla burgeri as its type species. genus: G. asterops and G. variabilis. Ghatixalus was The species of the subfamily range from being moder- morphologically diagnosed as follows: medium-sized ately large to small in size (SVL = 40–80 mm). Unlike adults (male SVL 38.8–51.3 mm, female 58.1–66.7 mm); other rhacophorids, their general body form appears a dorsal color pattern with dark brown prominent blotches; quite similar to that of . a free swimming tadpole stage, and the habitat associated The diagnosis of Buergeria is as follows: skin of body with mountain streams. Its distribution was restricted to and limbs smooth; no dorsal ornamentations; digital tips high altitudes in the of southern India (Biju moderately dilated and transverse groove sometimes et al., 2008; Frost, 2010). distinct; feet fully webbed; fingers up to half webbed Gracixalus was erected as a new subgenus of (Liem, 1970). The subfamily is distributed in Hainan and Aquixalus by Delorme et al. (2005) and its type species Taiwan (China), and in Japan from the Ryukyu Islands was Philautus gracilipes. Li et al. (2008) suggested to Honshu Island (Frost, 2010; Liem, 1970). Buergeria that Aquixalus (sensu stricto) should be placed within includes four species: B. buergeri, B. japonica, B. Kurixalus and Gracixalus was phylogenetically distant oxycephala, and B. robusta. from Kurixalus. Thus, Gracixalus was elevated to a full Chiromantis Peters, 1854, with type species genus. It was distinguished morphologically as follows: Chiromantis xerampelina, is the only Asian rhacophorid spines on the upper eyelid; rictal gland connected to genus that also occurs in Africa. Apart from Africa, the mouth; foot very thin; two outer palmar tubercles; white genus is distributed in East and Southeast Asia including spot on snout tip of tadpole, and five pairs of prelingual Myanmar, Thailand, Viet Nam, Laos and China (Frost, papillae and crescent shaped crest on the tadpole (Frost 2010). The species in this genus are moderately large et al., 2006). This genus, which ranges from southern with a SVL of 35–80 mm (Liem, 1970). Wilkinson et China southward to northwestern Thailand, central al. (2003) provided a key to distinguish between ten montane Viet Nam, and intervening Laos, has been species of Chirixalus. Asian species formerly assigned to considered to contain the following five species: G. Chirixalus are now assigned to Chiromantis to eliminate jinxiuensis, G. gracilipes, G. medogensis, G. quyeti and G. paraphyly (Frost et al., 2006; Grosjean et al., 2008; Li et supercornutus. al., 2008, 2009; Richards and Moore, 1998; Wiens et al., Kurixalus Ye, Fei and Dubois, 1999 has Rana eiffingeri 2009; Wilkinson et al., 2002). After Fei et al. (2010 b) as the type species. In the north, it is widely distributed in transferred Chiromantis vittatus into Feihyla, Chiromantis the Himalayan front ranges from eastern India to montane became monophyletic. Presently, Chiromantis cannot southern China and southward to southern Cambodia, be diagnosed morphologically due to the recent central Viet Nam, western and northern peninsular inclusion of the species formally assigned to Chirixalus. Thailand, northern Myanmar, Taiwan (China), and the Now, Chiromantis contains 15 species, including C. Ryukyu Islands of Japan (Frost, 2010). Compared with cherrapunjiae, C. doriae, C. dudhwaensis, C. hansenae, the species of Chiromantis, Kurixalus eiffingeri and C. kelleri, C. laevis, C. nongkhorensis, C. petersii, C. K. idiootocus do not have opposable phalanges and an punctatus, C. rufescens, C. samkosensis, C. senapatiensis, arch shaped anterior horn. In addition, vomerine teeth C. shyamrupus, C. simus and C. xerampelina (Fei et al., are present in K. eiffingeri and K. idiootocus. Kurixalus 2010b; Frost, 2010). presently includes ten species: K. ananjevae, K. Feihyla was erected by Frost et al. (2006) using baliogaster, K. banaensis, K. bisacculus, K. carinensis, Philautus palpebalis as the type species. This genus K. eiffingeri, K. idiootocus, K. naso, K. odontotarsus, occurs in southern Annam (Viet Nam) and Yunnan (China), and K. verrucosus (Frost, 2010; Yu et al., 2010). Using and it is expected to occur in Laos and elsewhere in two mtDNA genes, Yu et al. (2010) reconstructed the 6 Asian Herpetological Research Vol. 2

phylogeny of the K. odontotarsus species group, and Bossuyt et al., 2009; Biju et al., 2010; Delorme et al., suggested that K. hainanus was a junior synonym of K. 2005; Grosjean et al., 2008; Li et al., 2008, 2009). bisacculus. Polypedates Tschudi 1838, with type species P. A new genus, Liuixalus, with type species Philautus leucomystax, contains 23 species. The genus ranges romeri, was erected by Li et al. (2008) due to the across eastern India, Southeast Asia, Philippines, phylogenetic relationships of L. hainanus, L. ocellatus Indonesia, and Borneo (Frost, 2010). Due to similar and L. romeri (Frost, 2010; Li et al., 2008, 2009). No morphologies, the species of Polypedates are easily diagnostic morphological characters have been proposed confused with those of Rhacophorus. Liem (1970) for this genus, which is restricted to Hong Kong, Hainan splits the largely-sized rhacophorids into the genera and Guangxi (China). Polypedates and Rhacophorus. Channing (1989) agrees Nyctixalus Boulenger, 1882, with type species N. with this assignment and does not consider , is widely distributed in the Philippines, and Polypedates to be sister taxa. Jiang et al. (1987) Malaysia, Sumatra, Java, Borneo, and southern Viet Nam also consider these genera to be distinct. Rao et al. (Frost, 2010). Smith (1931) considers that Nyctixalus (2006) recognize Polypedates as a distinct genus and should be placed in Philautus; Dubois (1981) removed suggest it is paraphyletic with respect to Rhacophorus, it from the synonymy of Philautus. Recent molecular yet some studies recognize Rhacophorus only (e.g., studies suggest that Nyctixalus is the sister taxon of Dubois, 1987; Fei, 1999; Fei et al., 2005). Matsui and Theloderma and is distantly related to Philautus (Grosjean Wu (1994) cannot distinguish between Rhacophorus and et al., 2008; Li et al., 2008, 2009; Wilkinson et al., 2002; Polypedates on the basis of vocalizations. Molecular Yu et al., 2009). All the species of Nyctixalus share phylogenetic studies consistently support the recognition the following morphological characters: a medium- of both genera (Frost et al., 2006; Grosjean et al., 2008; sized adult being 30–40 mm in SVL; skin of head co- Li et al., 2008, 2009; Wilkinson et al., 2002; Yu et al., ossified to skull; body and limbs with numerous spiny 2008, 2009) and this assignment is widely accepted (Fei tubercles; fingers free or webbed at base; and vocal sacs et al., 2009, 2010a; Frost et al., 2006; Grosjean et al., always absent (Liem, 1970). It contains three species: N. 2008; Li et al., 2008, 2009; Wilkinson et al., 2002; Yu margaritifer, N. pictus, and N. spinosus. et al., 2008, 2009). Polypedates and Rhacophorus have Philautus Gistel, 1848, with type species Hyla several synapomorphic characters, including “Y”-shaped aurifasciata, includes 48 species. It ranges from terminal phalanges. Webbing between the fingers of central India through Myanmar and Thailand, south to Polypedates separates it from Rhacophorus (Liem, 1970; the Philippines and the Greater Sunda Islands (Frost, Wilkinson and Drewes, 2000; Rao et al., 2006). 2010). Previous morphological studies indicate that all Pseudophilautus Laurent, 1943 has Ixalus temporalis rhacophorids having aerial direct development of eggs as its type species. Li et al. (2009) resurrect the genus into froglets should be assigned into Philautus (Dring, from the synonymy of Philautus. They recognize the 1979; Bossuyt and Dubois, 2001). However, this form species within the Sunda shelf as Philautus sensu of development is not documented for most species and stricto, and the group with its centre of diversification the assignment to Philautus is usually based on overall on the Indian Subcontinent as Pseudophilautus. This morphological similarity alone. Dubois (1987) divides arrangement is consistent with Grosjean et al. (2008), Philautus into three subgenera: Philautus, Gorhixalus, Li et al. (2008), Meegaskumbura et al. (2002), and Yu and Kirtixalus. Bossuyt and Dubois (2001) revise this et al. (2008, 2009), and accepted by Frost (2010). Most taxonomy, transferring several Indian species from the recently, Biju et al. (2010) have erected the new genus subgenus Kirtixalus to Philautus. Recent molecular Raorchestes out of Pseudophilautus. Many new species studies suggest that Philautus is not monophyletic, and of Sri Lankan shrub-frogs are now known (Gururaja et some species are now transferred to other genera (Delorme al., 2007; Manamendra-Arachchi and Pethiyagoda, 2005; et al., 2005; Frost et al., 2006; Grosjean et al., 2008; Li Meegaskumbura and Manamendra-Arachchi, 2005), all et al., 2008, 2009; Meegaskumbura et al. 2002; Yu et al., of which are assigned to Pseudophilautus (Biju et al., 2008, 2009). Many Indian and Sri Lankan species are in 2010). Pseudophilautus (Li et al., 2009). Several new genera Raorchestes has Ixalus glandulosus as its type and species are now recognized and some subgenera are species (Biju et al. (2010). The genus is diagnosed by elevated to generic rank, such as Feihyla, Gracixalus, the following suite of characters: (1) relatively small Liuixalus, Pseudophilautus and Raorchestes (Biju and frogs, SVL of adult 1.5–4.5 cm; (2) vomerine teeth No. 1 Yaping ZAHANG et al. The Systematic Status of Rhacophorid Frogs 7 absent; large gular pouch transparent while calling; adults (SVL = 30–60 mm); numerous warts on body (3) primarily nocturnal; and (4) direct development and limbs, fingers webbed at base or up to half webbed. without free-swimming tadpoles (Biju et al., 2010). The inclusion of Theloderma moloch renders the genus According to the trees made by Li et al. (2009) and Yu paraphyletic and this species may represent a new genus et al. (2009), Chinese Pseudophilautus menglaensis and (Li et al., 2009). Notwithstanding, at present the genus P. longchuanensis, and Vietnamese P. gryllus should Theloderma contains 17 species: T. andersoni, T. asperum, be assigned to Raorchestes (Biju et al., 2010). Thus, T. baibengensis, T. bicolor, T. corticale, T. gordoni, T. Raorchestes ranges at least from the Western Ghats of horridum, T. kwangsiense, T. lateriticum, T. leporosum, India through mainland Southeast Asia to southern China, T. licin, T. moloch, T. nagalandense, T. phrynoderma, T. and the genus is not documented to occur in Sri Lanka rhododiscus, T. ryabovi and T. stellatum (Frost, 2010). (Biju et al., 2010). Containing 80 species, Rhacophorus Kuhl et Van 5. Suggestions for Future Research Hasselt, 1882 occurs widely across India, China, Japan, mainland Southeast Asia, the Greater Sunda Islands, and 5.1 Taxonomic representation Previous studies pro- the Philippines (Frost, 2010). The genus is diagnosed vide an initial resolution to the systematics and phy- by the following morphological characters: procoelus logeny of rhacophorids. A far more comprehensive vertebrae, anal folds that are usually present, typically investigation should be pursued and this should include extensive dermal forearm and tarsal folds (Bordoloi et as many species as possible. A densely-sampled tree is al., 2007; Jiang et al., 1987; Liem, 1970; Wilkinson and very important and necessary to gain an understanding Drewes, 2000). Dubois (1987) resurrects of the rhacophorids. Many Southeast Asian taxa remain Peters, 1867 as a subgenus and he subdivides the subgenus to be evaluated, including many species from the Phi- Rhacophorus into ten groups. However, this division is lippines, Borneo, and Malaysia. The same is true for not followed by many researchers (e.g., Harvey et al., Indian and Sri Lankan species. 2002). Iskandar and Colijn (2000) raise the subgenus A comprehensive phylogeny of Rhacophorus needs to Leptomantis to full generic rank. This genus now contains be reconstructed. Previous studies focus on species from R. appendiculatus, the species of Philautus in Dubois’ India, China and Viet Nam (Frost et al., 2006; Grosjean (1987), and the species of the subgenus Gorhixalus in et al., 2008; Li et al., 2008, 2009; Meegaskumbura et Bossuyt and Dubois’ (2001) (the Philautus hosii group of al., 2002; Richards and Moore, 1998; Wilkinson et al., Dring, 1987). However, Leptomantis cannot be diagnosed 2002; Yu et al., 2008, 2009). Most of the species from morphologically because its defining characters are shared Malaysia, mainland Southeast Asia and the Philippines are with some species of Rhacophorus sensu stricto, such not represented in existing data sets. Their phylogenetic as R. kajau, R. bisacculus and R. everetti (Dring, 1983; positions remain unknown. Inger et al., 1999; Liem, 1970). Recent molecular studies The phylogenetic placement of most putative species (Grosjean et al., 2008; Li et al., 2008, 2009; Wilkinson et of Philautus should be examined. Due to their small al., 2002; Yu et al., 2008, 2009) resolve two main groups size and cryptic habits, some species of Philautus are of Rhacophorus. Regardless, the validity of the subgenera represented in few studies. Many species are tentatively remains largely untested because previous studies do not assigned to Philautus based largely on their observed or include the type species L. bimaculata of the subgenus inferred direct developmental reproductive mode. Field Leptomantis. data are lacking for many species and thus, some former Theloderma Tschudi, 1838, with type species T. species of Philautus are now placed in other genera, such leporosa, is widely distributed in mainland Southeast as Kurixalus and Rhacophorus (Bossuyt and Dubois, Asia, including Sri Lanka, northeastern India, Myanmar 2001). Recent molecular studies suggest the existence of and southern China, and also Malaysia, and Sumatra much confusion. For example, many Chinese Philautus, (Frost, 2010). Frost et al. (2006) consider Theloderma to such as P. ocellatus and P. hainanus are now assigned to be the sister taxon of Philautus, but their analysis does Liuixalus. Philautus longchuanensis and P. gryllus are not include true Philautus. Li et al. (2008, 2009) and Yu now placed in Pseudophilautus (Grosjean et al., 2008; et al. (2009) resolve Theloderma as the sister taxon of Li et al., 2008, 2009). Further, Philautus banaensis now Nyctixalus; this clade forms the sister group of all other resides within Kurixalus, and P. quyeti within Gracixalus rhacophorids excluding Liuixalus and Buergeria. The (Grosjean et al., 2008; Li et al., 2008, 2009). The new diagnostic characters are as follows: medium to large genera Pseudophilautus and Raorchestes, have close 8 Asian Herpetological Research Vol. 2 phylogenetic relationships with Philautus, to the extent tory typical of other taxa. Previous studies (Grosjean et that morphological diagnosis remains problematic. Future al., 2008; Li et al., 2008, 2009; Meegaskumbura et al., studies should focus on the phylogeny and taxonomy of 2002; Yu et al., 2008, 2009) suggest that multiple taxa the complicated genera Philautus, Pseudophilautus and have their origins on the Indian subcontinent, and that Raorchestes. some genera disperse out of Indian multiple times to East and Southeast Asia. Some may have returned back 5.2 Discovery of morphologically diagnostic characters to India. Further investigation is highly desirable. A robust molecular phylogeny should be used to identify potential morphological characters that can serve to Acknowledgements We thank Darrel FROST, Jinzhong diagnose higher taxa. Future studies should also focus on FU and Shengxian ZHONG for their critical reviews and the discovery of morphological characters that can lead discussions on the manuscript. This work was supported to the construction of a total evidence morphological and by China Postdoctoral Science Foundation Funded molecular analysis. Project (No. 20100471689), the ‘Xibuzhiguang’ Project 5.3 The study of character evolution Given a robust of Chinese Academy of Sciences, the ‘Lingyuqianyan’ molecular phylogeny, future research can center on the Project of Chinese Academy of Sciences to Jiatang mechanisms of adaptation to the surrounding environ- LI, the National Natural Science Foundation of China ment. Extensive adaptive radiations involving parallel (Grant Nos. 30911120031, 31071894, 30670243) and and convergent evolution occur in the Caudata and the Basic Research Program of the National Science and Anura (e.g., Wake, 1966, 1987; Zhang et al., 2006). Technology of China (Grant Nos. 2006FY110500-3, Bossuyt and Milinkovitch (2000) provide a nice ex- 2008GA001). ample of what is possible to achieve. Future research on morphological character evolution will be valuable. References For example, the evolution of toe webbing is a remar- kable feature in rhacophorids and it may represent Ahl E. 1931. Amphibia Anura III. Polypedatidae. Das Tierreich, Berlin: Lieferung, 55, xvi, 1–477 adaptive parallelism within different lineages. Amphibia Web. 2010. Information on amphibian biology and 5.4 Biogeography of widely distributed species Most conservation [web application]. Available at http://amphibiaweb. rhacophorids live on trees, but some inhabit smaller org (accessed 2010) Bahir M. 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