Triprion Cope, 1866

FAIVOVICH, JULIÁN, HADDAD, CÉLIO F. B., GARCIA, PAULO C. A., FROST, DARREL R., CAMPBELL, JONATHAN A. & WHEELER, WARD C., 2005, Systematic Review Of The Frog Family Hylidae, With Special Reference To Hylinae: Phylogenetic Analysis And Taxonomic Revision, Bulletin of the American Museum of Natural History 2005 (294), pp. 1-240 : 107-108

Triprion Cope, 1866

TYPE SPECIES: Pharyngodon petasatus

Cope, 1865, by monotypy.

Pharyngodon Cope, 1865 . Junior homonym of Pharyngodon Diesing, 1861 . Type species: Pharyngodon petasatus Cope, 1865 , by monotypy.

Diaglena Cope, 1887 . Type species: Triprion spatulatus Günther, 1882 , by monotypy.

DIAGNOSIS: For the purposes of this paper we consider that the 125 transformations in nuclear and mitochondrial protein and ribosomal genes autapomorphic of Triprion petasatus are synapomorphies of this genus. See appendix 5 for a complete list of these molecular synapomorphies. In Duellman’s (2001) phylogenetic analysis of Pternohyla , Smilisca , and Triprion , the monophyly of Triprion is supported by three synapomorphies 28: maxilla greatly expanded laterally; prenasal bone present (known homoplastic instance in Aparasphenodon ); and presence of parasphenoid odontoids.

COMMENTS: We included a single species of this genus, and as such we did not test its monophyly, but we do not consider it controversial on the basis of the morphological evidence mentioned above.

CONTENTS: Two species. Triprion petasatus (Cope, 1865) ; Triprion spatulatus Günther, 1882 .

LOPHIOHYLINI MIRANDA­ RIBEIRO, 1926

Lophiohylinae Miranda­Ribeiro, 1926. Type genus: Lophyohyla Miranda­Ribeiro, 1926 .

Trachycephalinae B. Lutz, 1969. Type genus: Trachycephalus Tschudi, 1838 .

DIAGNOSIS: This tribe is diagnosed by 63 transformations in nuclear and mitochondrial protein and ribosomal genes. See appendix 5 for a complete list of these molecular synapomorphies. A putative morphological synapomorphy of this tribe is the presence of at least four posterior labial tooth rows in the larval oral disc (e.g., Bokermann, 1966b; Duellman, 1974; de Sa´, 1983; Lannoo et al., 1987; McDiarmid and Altig, 1990; Schiesari et al., 1996; da Silva in Altig and Mc­ Diarmid, 1999b; Wogel et al., 2000) (reversals in Osteopilus marianae , O. crucialis , O. wilderi [Dunn, 1926] and in Osteocephalus oophagus [Jungfer and Schiesari, 1995]).

COMMENTS: This tribe contains all South American and West Indian casque­headed frogs and related groups. It includes Aparasphenodon , Argenteohyla , Corythomantis , Osteopilus , Phyllodytes , Tepuihyla , a new monotypic genus, and the genera Osteocephalus and Trachycephalus as redefined here.

Recently, Kasahara et al. (2003) noticed that Aparasphenodon brunoi , Corythomantis

28 Note that on his preferred tree (fig. 410) one of these character transformations is numbered 18, which seems to be a typographical error for 12, the only other character that supports this clade but that is not shown in the tree.

greeningi , and Osteocephalus langsdorffii share similar chromosome morphology, where there is a clear discontinuity in the chromosome lengths of the first five pairs and the remaining seven pairs. Furthermore, they share the presence of a secondary constriction in pair 10. Available information on karyotypes of other casque­headed frogs of this clade suggests that the discontinuity in chromosome lengths occurs as well in Argenteohyla (apparent from plates published by Morand and Hernando, 1996), Phrynohyas venulosa (apparent from plates published by Bogart, 1973), and some species of Osteopilus ( O. brunneus , O. dominicensis , O. marianae , O. septentrionalis ), but not in Osteocephalus taurinus , the only species of the genus Osteocephalus , as redefined here, whose karyotype was studied (Anderson, 1996). The position of the secondary constriction also varies, having been observed in chromosome 4 in Argenteohyla (Morand and Hernando, 1996) , chromosome 9 in Osteopilus brunneus , O. dominicensis , O. septentrionalis , and O. wilderi (Anderson, 1996) , chromosome 10 in Phrynohyas venulosa (apparent from plates published by Bogart, 1973), and in chromosome 12 in Osteocephalus taurinus . The taxonomic distribution of these character states needs further study to define the inclusiveness of the clades they support.

Delfino et al. (2002) noticed that serous skin glands of Osteopilus septentrionalis and Phrynohyas venulosa produce secretory granules with a dense cortex and a pale medulla; they observed the same in a photograph of a section of skin of Corythomantis greeningi published by Toledo and Jared (1995). Very few hylid taxa were studied for serous gland histology, and these include a few species of Phyllomedusa, Holarctic Hyla , Scinax , and Pseudis paradoxa (see Delfino et al., 2001, 2002). The taxonomic distribution of these peculiar secretory granules requires additional study to assess its level of generality and the clade or clades that it diagnoses.