Rana labialis, BOULENGER, 1887

Inger, Robert F., Stuart, Bryan L. & Iskandar, Djoko T., 2009, Systematics of a widespread Southeast Asian frog, Rana chalconota (Amphibia: Anura: Ranidae), Zoological Journal of the Linnean Society 155 (1), pp. 123-147 : 130-132

RANA LABIALIS BOULENGER

(Previously referred to as Peninsular Malaysia morphotype)

Rana labialis Boulenger, 1887: 345 – Malacca; van Kampen, 1923: 220; Stuart et al., 2006: 473.

Student’s t -test used for SVL, Mann–Whitney test used for body proportions. Abbreviations are as defined in the Material and Methods.

Rana chalconota (part) Boulenger, 1920: 201; Smith, 1930: 109.

Rana chalconota raniceps (part) Inger, 1966: 177.

The type locality of this taxon is Malacca, Peninsular Malaysia. Boulenger (1887) stated that the types of labialis had dark spots dorsally, illustrated in his plate X, fig. 1, and that males had no humeral glands. Boulenger (1920) gave the SVL range of three male syntypes as 34–36 mm and that of three female syntypes as 42–49 mm. We have examined the type series and have detected weak humeral glands in the

*Female FRIM 1539, male FRIM 829.

†Female FRIM 1418, male FRIM 826; = unnamed lineage of Figure 2 .

‡Females FRIM 1539, 1048, males FRIM 1047, 1415; tissues not available.

§+ = back spotted, 0 = back not spotted.

males, which have constricted nuptial pads. Our measurements are 33.6–35.8 mm for two syntypic males and 48.8 mm for a single female. We here designate that female, BMNH 1947.2 .3.40, as the lectotype of Rana labialis Boulenger. A description of the lectotype is given below .

The relationships and specific assignments of frogs from Peninsular Malaysia are not clear. The continental populations ( Thailand and Peninsular Malaysia) of the chalconota -like frogs clearly exhibit a great deal of genetic structure. The molecular phylogenetic analysis ( Fig. 2 ) shows three distinct, genetically divergent clades (maximum uncorrected pairwise sequence divergence of 11.90%; Table 5) from Peninsular Malaysia, one of which is phylogenetically unrelated to the other two ( Fig. 2 ). We attempted to amplify historical mtDNA from the lectotype of labialis following the extraction protocol of Stuart et al. (2006), but did not succeed. Consequently, assigning any of these clades to labialis must at this point depend on morphological data alone.

The Peninsular Malaysian specimens (and tissues) are from three localities: Gunung Jerai (about 450 km from the type locality of labialis ), Penang Island (about 410 km) and Selangor (about 125 km). Five specimens from Penang Island and three from Gunung Jerai, in northern Peninsular Malaysia, constitute a clade (‘ cf. labialis ’; Fig. 2 ), ten from Selangor (‘ labialis ’; Fig. 2 ) constitute another and four more from Gn. Jerai form the third, distantly related clade (‘unnamed lineage’; Fig. 2 ). We have examined an additional 22 long-preserved specimens from Selangor without associated tissues. All Selangor specimens resemble the lectotype and the other type specimens of labialis in the prevalence of dorsal spotting; all of the types have the dorsal spots and 25 of the 32 from Selangor are also spotted. The Selangor frogs also agree with types in size and body proportions ( Table 7), but differ from frogs from nearby southern Thailand ( Table 8). We assign these Selangor frogs to the species labialis .

Taxonomic assignment of the ‘ cf. labialis ’ Malaysian frogs is not certain. They differ from the types of labialis and the Selangor frogs we assign to that species in dorsal spotting; only one of seven ‘ cf. labialis ’ frogs is spotted. Males of this clade have slightly wider heads (median 0.323 of SVL) than males from Selangor (median 0.296); the two arrays differ significantly (Mann–Whitney test; P <0.02). Yet the morphological differences between these two Malaysian clades are not as great as the differences between sympatric pairs of species, e.g. raniceps and megalonesa in Borneo (see

Figures

Figure 2. Fifty per cent majority-rule consensus phylogram resulting from mixed-model Bayesian analysis of mitochondrial DNA from frogs of the Rana chalconota species group. Trees were rooted with R. erythraea and R. nigrovittata (not shown). Numbers above and below nodes are Bayesian posterior probabilities and parsimony bootstrap values> 50, respectively. An asterisk (*) indicates a Bayesian posterior probability of 1.00 and parsimony bootstrap value of 100. Maximum parsimony analysis resolved the clades Peninsular Malaysia D [Peninsular Malaysia C (Thailand + Padang Small)], but without bootstrap support.