Evolution of reproduction in the Rhacophoridae (Amphibia, Anura)

Abstract

Rhacophorid treefrogs have different reproductive modes: some go through a tadpole stage and some have direct development, and the adults of some species produce foam nests. Philautus is the only genus characterized by direct development. The production of foam nests has been reported in the genera Polypedates, Rhacophorus, Chiromantis and Chirixalus. Recent molecular studies did not provide a robust hypothesis concerning the origin of these reproductive modes in the Rhacophoridae. In order to better understand the evolution of these reproductive modes, we tried to clarify relationships within this group, using DNA sequencing. Our data set consists in a large number of new sequences (1676 base pairs corresponding to threee genes) for five outgroup ranoids and 48 Rhacophoridae, including 16 undescribed species from Sri Lanka and southern India, and all homologous data available in Genbank. After the inclusion of Philautus from India, our data show that the separation of Philautus into clades does not coincide with their geographic distribution. Our data point to the existence of a clade, including the genera Rhacophorus, Polypedates, Chiromantis and Chirixalus, which confirms the results of Wilkinson et al. (2002) and suggests that the ability to produce foam nests has emerged only once in the Rhacophoridae, as already stated by these authors.

The Systematic Status of Rhacophorid Frogs

“…Marmayou et al (2000) suggested that rhacophorids were deeply imbedded within the Ranidae. In their anatomical and molecular analyses, Frost et al (2006) recognized the family Rhacophoridae and this usage was consistent with most other studies (Bossuyt and Roelants, 2009;Fei et al, 2009;Grosjean et al, 2008;Jiang et al, 1987;Li et al, 2008Li et al, , 2009Yu et al, 2008Yu et al, , 2009Zhao and Adler, 1993;Zhao et al, 2000). Channing's recognition of the two subfamilies has been strongly supported by several molecular studies (Bossuyt et al, 2006;Frost et al, 2006;Li et al, 2008Li et al, , 2009Wilkinson et al, 2002;Yu et al, 2009).…”

“…The clades of Philautus did not coincide with geographic distributions. Grosjean et al (2008) also studied the evolution of reproduction in the Rhacophoridae and found that the ability to produce a foam nest emerged only once, as previously proposed (Wilkinson et al, 2002). Li et al (2008) employed maximum parsimony, maximum likelihood, and Bayesian inference analyses to investigate the molecular phylogenetic relationships of the Rhacophoridae.…”

“…Using 1676 bp of mitochondrial and nuclear gene fragments (12S, 16S and rhodopsin) from 48 rhacophorids, including 16 undescribed Sri Lankan and southern Indian species, Grosjean et al (2008) recognized both subfamilies Buergeriinae and Rhacophorinae and erected the new tribe Nyctixalini. The basal position of Buergeria within the Rhacophorinae was well supported, and Nyctixalus and Theloderma were sister taxa.…”

“…The Philautini contained "Aquixalus", Kurixalus, Nyctixalus, Philautus, and Theloderma, and the Rhacophorini included Chirixalus, Chiromantis, Polypedates, and Rhacophorus (Dubois, 2005). Using mitochondrial DNA (mtDNA) and nuclear DNA (nuDNA) sequence data, Grosjean et al (2008) suggested that the Rhacophorinae should consist of two tribes, the Rhacophorini and Nyctixalini, the latter containing Theloderma and Nyctixalus. Using an even larger DNA sequence dataset, Li et al (2009) Buergeriinae has been maintained without modification (Channing, 1989;Delorme et al, 2005;Frost et al, 2006;Grosjean et al, 2008;Li et al, 2008Li et al, , 2009Liem, 1970;Wilkinson et al, 2002;Yu et al, 2009).…”

Eight new species of Pseudophilautus (Amphibia: Anura: Rhacophoridae) from Sripada World Heritage Site (Peak Wilderness), a local amphibian hotspot in Sri Lanka

“…These direct developing rhacophorids (Bossuyt & Dubois 2001;Meegaskumbura et al 2002;Bahir et al 2005;Grosjean et al 2008;Li et al 2008Li et al , 2009Biju & Bossuyt 2009;Yu et al 2009) previously belonged to the genus Philautus which were widely distributed throughout tropical Asia (Bossuyt & Dubois 2001;Bossuyt et al 2004;Frost et al 2006;Yu et al 2009). But later, considering molecular phylogenetics, the populations in Sri Lanka and India were found to be a separate radiation belonging to a separate genus known now as Pseudophilautus (Li et al 2009;Yu et al 2010;Meegaskumbura & ManamendraArachchi 2011).…”