AENIGMANURA, Brown & Siler & Richards & Diesmos & Cannatella, 2015

Brown, Rafe M., Siler, Cameron D., Richards, Stephen J., Diesmos, Arvin C. & Cannatella, David C., 2015, Multilocus phylogeny and a new classification for Southeast Asian and Melanesian forest frogs (family Ceratobatrachidae), Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 174 (1), pp. 130-168 : 152-153

AENIGMANURA SUBGEN. NOV.

Type species

Platymantis papuensis schmidti Brown & Tyler, 1968 .

Diagnosis

Individual species of the subgenus Aenigmanura differ from other members of Cornufer by characters related to their general classification as either generalized terrestrial species with narrow finger and toe discs or arboreal forms with widely expanded finger and toe discs. The arboreal tree frogs of Aenigmanura ( Cornufer citrinospilus, Co. custos , Co. hedigeri [formerly Platymantis guppyi ; see below], Co. macrosceles, Co. mamusiorum, Co. nakanaiorum , Co. neckeri , Co. nexipus, Co. parilis, Co. sp. B. Manus and Co. sp. C. Manus) can be readily distinguished from ground frogs (selected members of subgenus Aenigmanura and all Cornufer species not assigned to subgenera, all with non-expanded discs of fingers and toes), aquatic species (subgenera Potamorana and Discodeles , characterized by the presence of interdigital webbing), sticky-toed frogs (subgenus Batrachylodes , small, triangularshaped bodies with strongly pointed snouts), the Fijian ground frog ( Co. vitianus , with non-expanded discs of fingers and toes), palm frogs (subgenus Palmatorappia , with interdigital webbing present on hands), and horned frogs (subgenus Ceratobatrachus , with elaborately casqued skull morphology and dermal horns above the eyelids). The terrestrial species of Aenigmanura ( Cornufer adiastolus , Cornufer admiraltiensis , Cornufer akarithymus , Cornufer boulengeri , Cornufer bufonulus , Cornufer desticans , Cornufer gilliardi , Cornufer latro , Cornufer magnus , Cornufer papuensis , Cornufer pelewensis , Cornufer schmidti , Cornufer solomonis , Cornufer sulcatus , and Cornufer weberi ) can be distinguished from river frogs and giant frogs (subgenera Potamorana and Discodeles , characterized by the presence of interdigital webbing), sticky-toed frogs (subgenus Batrachylodes , small, triangular-shaped bodies with strongly pointed snouts), the Fijian tree frog ( Co. vitiensis , with widely expanded discs of fingers and toes), palm frogs (subgenus Palmatorappia , an arboreal species with expanded finger and toe discs, and interdigital webbing present on hands and feet), and horned frogs (subgenus Ceratobatrachus , with elaborately casqued skull morphology and dermal horns above the eyelids).

As implied by the name, the wide range of morphological and ecological variation in this clade renders an unambiguously exclusive diagnosis of Aenigmanura impossible. We are unaware of morphological synapomorphies for this group, although our phylogenetic analysis provides very strong support for this phenotypically and ecologically diverse clade ( Fig. 2, Clade J).

Phylogenetic definition

Aenigmanura (NCN) is a maximum crown-clade name referring to the crown clade ( Fig. 2, Clade J) originating with the most recent common ancestor of Co. papuensis and all extant species that share a more recent common ancestor with Co. papuensis than with any of the other species of the clade Cornufer . Alternatively it can be conceived of as the largest crown clade containing Co. papuensis , but not any other species of the clade Cornufer .

Content

Cornufer adiastolus, Co. admiraltiensis , Co. akarithymus , Co. boulengeri , Co. citrinospilus, Co. custos, Co. desticans , Co. gilliardi , Co. hedigeri (formerly Pl. guppyi ; see below), Co. latro , Co. macrosceles, Co. magnus, Co. mamusiorum, Co. nakanaiorum , Co. neckeri , Co. nexipus, Co. parilis , Co. papuensis , Co. pelewensis, Co. schmidti , Co. solomonis, Co. sulcatus , Co. weberi , the newly described Co. custos ( Richards et al., 2014) , and two undescribed species from Manus Island (sp. B Manus and sp. C Manus; Fig. 2., Clade J).

Comment

The range of body sizes in this large clade is striking. From miniaturized terrestrial species such as Co. sulcatus and Co. akarithymus (males 17–27 mm SVL), to giant ground species such as Co. magnus (males 75–150 mm SVL), to large canopy frogs such as Co. neckeri and Co. hedigeri (formerly Pl. guppyi ; see below), to delicate, high-elevation, arboreal shrub species such as Co. macrosceles and Co. mamusiorum , to widespread terrestrial generalists such as Co. papuensis , Co. weberi, Co. schmidti , and Co. solomonis – the subgenus Aenigmanura exhibits nearly the full range of ceratobatrachid ecomorphological diversity ( Brown, 2004), all within one clade of closely related species.

As noted above, allocation of Pl. guppyi Boulenger, 1884 (not to be confused with D. guppyi ) to the genus Cornufer presented a case of secondary homonymy with respect to R. guppyi Boulenger, 1884 . Given that no available junior synonym exists for the latter and that it is also the type species of Discodeles , we elected not to alter this name, and we have given R. guppyi precedence over Pl. guppyi following the principle of first reviser (ICZN, 1999:30). Thus, the establishment of a replacement name for Pl. guppyi Boulenger, 1884 , is necessary. Accordingly, we designate Co. hedigeri as a nomen novum for Pl. guppyi Boulenger, 1884 . The epithet hedigeri is a patronym for Heine Hediger (1908–1992) in recognition of his contributions ( Hediger, 1933, 1934) to the taxonomy of the genus Cornufer sensu lato and the biology of the South Pacific.

Etymology

From the Latin enigma, meaning something ‘obscure or unknown, a riddle’, and anura, meaning ‘frog’, in reference to the unanticipated and confusing range of morphological and ecological variation represented by the closely related species of the new subgenus. The name is masculine in gender. Suggested common name: Pacific Island riddle frogs.