Phylogenetic relationships and generic taxonomy of the tribe Paini (Amphibia, Anura, Ranidae, Dicroglossinae), with diagnoses of two new genera Annemarie OHLER Alain DUBOIS Muséum national d’Histoire naturelle, Département Systématique et Évolution, Reptiles et Amphibiens, USM 0602, case postale 30, 25 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Ohler A. & Dubois A. 2006. — Phylogenetic relationships and generic taxonomy of the tribe Paini (Amphibia, Anura, Ranidae, Dicroglossinae), with diagnoses of two new genera. Zoosystema 28 (3) : 769-784. ABSTRACT KEY WORDS A preliminary cladistic analysis of the relationships between 26 frog species Amphibia, of the tribe Paini (Ranidae, Dicroglossinae) was carried out on the basis of 31 Anura, Ranidae, morphological characters, mainly from external morphology of adults. Combined Dicroglossinae, with the results of a molecular analysis published elsewhere, these data 1) confi rm Paini, that, after exclusion of the species Rana delacouri Angel, 1928, the Paini are a morphology, cladistic relationships, homophyletic group, and 2) allow to redefi ne the genera of this tribe, which new genera. are now six in number, including two new ones described herein. RÉSUMÉ Relations phylogénétiques et taxonomie générique de la tribu Paini (Amphibia, Anura, Ranidae, Dicroglossinae), avec diagnoses de deux nouveaux genres. Une analyse cladistique préliminaire des relations entre 26 espèces de grenouilles MOTS CLÉS de la tribu des Paini (Ranidae, Dicroglossinae) a été menée en utilisant 31 Amphibia, caractères morphologiques, principalement de la morphologie externe des Anura, Ranidae, adultes. Combinées avec les résultats d’une analyse moléculaire publiée ailleurs, Dicroglossinae, ces données 1) confi rment qu’après exclusion de l’espèce Rana delacouri Angel, Paini, 1928, les Paini constituent un groupe homophylétique, et 2) permettent de morphologie, analyse cladistique, redéfi nir les genres de cette tribu, qui sont maintenant au nombre de six, dont genres nouveaux. deux genres nouveaux décrits ici. ZOOSYSTEMA • 2006 • 28 (3) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 769 Ohler A. & Dubois A. INTRODUCTION lection; Museum of Comparative Zoology, Harvard (MCZ); Muséum national d’Histoire naturelle, We recently (Dubois & Ohler 2005) described the Paris (MNHN); Naturhistorisches Museum, Wien male secondary sex characters of Chaparana aenea (NMW); Senckenberg Museum, Frankfurt (SMF); (Smith, 1922), which had remained unknown since Zoological Museum Kobenhavn (ZMK). Appendix the discovery of the species. Th ese observations presents a list of the specimens examined for this suggest that the provisional taxonomy proposed analysis with their localities. In the text below, in by Dubois (1992) for this frog and related species Figure 1 and Table 1, we use the specifi c names (distributed in two genera, Chaparana Bourret, of the species studied “naked”, without generic 1939 and Paa Dubois, 1975, grouped in a tribe names, pending the taxonomic proposals of our Paini) cannot be maintained, and lead us to suggest conclusion. some changes in this scheme. Th e discovery of the Specimens were sexed either according to their sexual secondary characters of this species implies external characters (in the case of adult breeding taxonomic changes for the whole group where it males) or through a slight lateral incision in order belongs. We provide below a fi rst tentative cladistic to examine the gonad: the state of development of analysis of this group of frogs based on external the latter and its ducts allowed to distinguish two morphological characters. Following the results developmental stages, juvenile (including subadult) of the work of Jiang & Zhou (2005), the genus and adult, as explained in detail by Dubois (1976: Nanorana Günther, 1896 was included in this 31-33). A specimen was radiographed using a analysis, and the tribe Dicroglossini (as defi ned Hewlett-Packard 43805N X-Ray System apparatus by Dubois 2003) was considered the sister-taxon (Faxitron Series). of the Paini, in the subfamily Dicroglossinae of the family Ranidae. Combined with the results CHARACTERS AND CHARACTER STATES of molecular studies published elsewhere (Jiang In order to be able to use as many species as possible, et al. 2005), these results allow to propose a new including rare ones or species for which we had taxonomy for this tribe, with diagnoses of two access to a single or a few specimens, we mainly new genera. studied characters of the external morphology, that can be scored without damaging the specimens. We scored 0 the characters states as observed in the MATERIAL AND METHODS outgroups but no polarity of character states was given prior to analysis. Table 1 gives the distribution MATERIAL of character states in the species studied, according For this analysis, the ingroup was composed of 25 to the following defi nitions of character states: species of the genera Chaparana and Paa as defi ned (C01) Warts on mid-dorsal skin: [0] absent or by Dubois (1992) and one species of the genus indistinct; [1] present. Nanorana. For the outgroup, given the results of (C02) Aspects of warts on mid-dorsal skin: [0] the studies of Jiang & Zhou (2005) and Jiang et al. rounded or slightly elongate, but if elongate not (2005), we used four species of the dicroglossine regularly arranged on back; [1] longitudinally genera Euphlyctis Fitzinger, 1843, Fejervarya Bolkay, elongate, regularly arranged on back. 1915, Hoplobatrachus Peters, 1863 and Limnonectes (C03) Latero-dorsal folds: [0] absent; [1] discontinu- Fitzinger, 1843, as defi ned by Dubois (1992, 2003). ous or replaced by warts more or less in line; [2] Specimens studied are from the following collections: narrow and continuous all along back. American Museum of Natural History, New York (C04) Tympanum: [0] distinct; [1] indistinct in (AMNH); Natural History Museum, London external examination. (BMNH); Chengdu Institute of Biology, Academia (C05) Relative length of fi ngers I and II: [0] fi nger Sinica, Chengdu, Sichuan (CIB); Malcolm A. Smith I longer than II; [1] fi nger I shorter than or as collection (MAS), now housed in the BMNH col- long as II. 770 ZOOSYSTEMA • 2006 • 28 (3) Phylogeny and taxonomy of the Paini (Amphibia, Anura) (C06) Tips of toes: [0] pointed or blunt; [1] en- if applicable) of adult male in breeding condition: larged. [0] small, indistinct, uncountable; [1] large, distinct, (C07) Proximal subarticular tubercles of fi ngers: countable. [0] small or medium; [1] large. (C24) Colour of nuptial spines on fi ngers and breast (C08) Leg length: [0] shorter than half of snout-vent of adult male in breeding condition: [0] black or length; [1] longer than half of snout-vent length. brownish; [1] translucent or creamy. (C09) Webbing: [0] very incurved between extremi- (C25) Patches of nuptial spines on breast of adult male ties of adjacent toes; [1] slightly incurved between in breeding condition: [0] two well delimited patches extremities of adjacent toes; [2] complete, not of densely packed spines; [1] two separated patches incurved between extremities of adjacent toes. of unequally spaced spines; [2] two confl uent patches (C10) Flap of skin along toe V: [0] from tip of toe or a single patch covering both sides of chest. to level of fi rst subarticular tubercle of toe V; [1] (C26) Arrangement of spines on arms, breast and from tip of toe to between subarticular tubercle of belly of adult male in breeding condition: [0] toe V and base of metatarsus; [2] from tip of toe to isolated; [1] in clusters. base of metatarsus of toe V or nearly so. (C27) Vent of adult male in breeding condition: (C11) Tarsal fold: [0] present, well developed; [1] [0] without morphological diff erentiation; [1] with very weak or absent. a square dermal fl ap; [2] with spines around and (C12) Mid-dorsal chevron: [0] always absent; [1] inside vent. may be present. (C28) Intersexuality (adult females with nuptial spines (C13) Mid-dorsal line: [0] always absent; [1] may on fi nger I): [0] always absent; [1] may be present. be present. (C29) Colour of eggs: [0] with coloured animal (C14) Vocal sacs in adult male: [0] present; [1] pole; [1] entirely whitish or creamy. absent. (C30) Number of ridges bearing keratodont rows (C15) Forearm in adult male in breeding condition: on upper lip of tadpoles: [0] 1-3 rows, median [0] not enlarged; [1] enlarged. value in species ranges from 1.5 to 2 rows; [1] 2-5 (C16) Nuptial spines on prepollex and fi nger I of rows, median value in species ranges from 3 to 3.5 adult male in breeding condition: [0] present; [1] rows; [2] 4-8 rows, median value in species ranges absent. from 4.5 to 7 rows; [3] 7-9 rows, median value (C17) Nuptial spines on fi nger II of adult male in species ranges from 8 to 8.5 rows; [4] 0-1 row, in breeding condition: [0] always absent; [1] may median value in species 0.5 row. be present. (C31) Number of ridges bearing keratodont rows (C18) Nuptial spines on fi nger III of adult male on lower lip of tadpoles: [0] 2-3 rows, median value in breeding condition: [0] always absent; [1] may in species ranges from 2.5 to 3 rows; [1] 0-3 rows, be present. median value in species ranges from 1.5 to 2 rows; (C19) Nuptial spines on arm and forearm of adult [2] 3-5 rows, median value in species 4 rows. male in breeding condition: [0] always absent; [1] may be present. CLADISTIC METHODOLOGY (C20) Nuptial spines on anterior part of throat of We carried out a cladistic analysis based on the 31 adult male in breeding condition: [0] always absent; characters above in 30 species (Table 1), using PAUP, [1] may be present. version 4.0b10 (Swoff ord 2001). We conducted (C21) Nuptial spines on breast of adult male in heuristic searches with initial trees obtained from breeding condition: [0] always absent; [1] may random addition sequence using 100 replicates, be present.