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The Systematic Status of Rhacophorid Frogs

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The Systematic Status of Rhacophorid Frogs Asian Herpetological Research 2011, 2(1): 1-11 DOI: 10.3724/SP.J.1245.2011.00001 The Systematic Status of Rhacophorid Frogs 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 Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada Abstract The frog family Rhacophoridae recently underwent notable reorganization and taxonomic rearrangement. Several new genera were erected to reflect evolutionary history, including Ghatixalus, Gracixalus, Feihyla, Liuixalus and Raorchestes. 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, amphibian, 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 species in rhacophorids. Several genera, including Philautus, in 14 genera (Biju et al., 2010; Frost, 2010). The group Pseudophilautus 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, Myanmar, 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 Chiromantis, and southward across Indochina to Greater Sunda Feihyla, Rhacophorus and Polypedates 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 taxonomy 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 “Chirixalus”. Philautus rooted as the sister group to this the family Rhacophoridae, recognized 12 genera and clade and Buergeria being the most basal genus. 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 Nyctixalus 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 Rhacophorinae 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. Theloderma 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) Kurixalus was closely close relationship, and Rhacophorus formed the sister related to Aquixalus; and (3) Kurixalus idiootocus and K. clade to Philautus
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