A review of the systematics of the frog family Hylidae, with special reference to the Hylinae: a phylogenetic analysis and taxonomic revision
FAIVOVICH, J., HADDAD, C. F. B., GARCIA, P. C. A., FROST, D. R., CAMPBELL, J. A., WHEELER, W. C.
Bulletin of the American Museum of Natural History
American MUseum of Natiural History
Año: 2005 p. 1 - 1
Hylidae is a large family of American, Australopapuan, and temperate Eurasian treefrogsof approximately 870 known species, divided among four subfamilies. Although some groupsof Hylidae have been addressed phylogenetically, a comprehensive phylogenetic analysis hasnever been presented.The first goal of this paper is to review the current state of hylid systematics. We focus onthe very large subfamily Hylinae (590 species), evaluate the monophyly of named taxa, andexamine the evidential basis of the existing taxonomy. The second objective is to perform aphylogenetic analysis using mostly DNA sequence data in order to (1) test the monophyly ofthe Hylidae; (2) determine its constituent taxa, with special attention to the genera and speciesgroups which form the subfamily Hylinae, and c) propose a new, monophyletic taxonomyconsistent with the hypothesized relationships.We present a phylogenetic analysis of hylid frogs based on 276 terminals, including 228hylids and 48 outgroup taxa. Included are exemplars of all but 1 of the 41 genera of Hylidae(of all four nominal subfamilies) and 39 of the 41 currently recognized species groups of thespecies-rich genus Hyla. The included taxa allowed us to test the monophyly of 24 of the 35nonmonotypic genera and 25 species groups of Hyla. The phylogenetic analysis includesapproximately 5100 base pairs from four mitochondrial (12S, tRNA valine, 16S, and cytochromeb) and five nuclear genes (rhodopsin, tyrosinase, RAG-1, seventh in absentia, and28S), and a small data set from foot musculature.Concurring with previous studies, the present analysis indicates that Hemiphractinae are notrelated to the other three hylid subfamilies. It is therefore removed from the family and tentativelyconsidered a subfamily of the paraphyletic Leptodactylidae. Hylidae is now restrictedto Hylinae, Pelodryadinae, and Phyllomedusinae. Our results support a sister-group relationshipbetween Pelodryadinae and Phyllomedusinae, which together form the sister taxon ofHylinae. Agalychnis, Phyllomedusa, Litoria, Hyla, Osteocephalus, Phrynohyas, Ptychohyla,Scinax, Smilisca, and Trachycephalus are not monophyletic. Within Hyla, the H. albomarginata,H. albopunctata, H. arborea, H. boans, H. cinerea, H. eximia, H. geographica, H. granosa,H. microcephala, H. miotympanum, H. tuberculosa, and H. versicolor groups are alsodemonstrably nonmonophyletic. Hylinae is composed of four major clades. The first of theseincludes the Andean stream-breeding Hyla, Aplastodiscus, all Gladiator Frogs, and a Tepuianclade. The second clade is composed of the 30-chromosome Hyla, Lysapsus, Pseudis, Scarthyla,Scinax (including the H. uruguaya group), Sphaenorhynchus, and Xenohyla. The thirdmajor clade is composed of Nyctimantis, Phrynohyas, Phyllodytes, and all South American/West Indian casque-headed frogs: Aparasphenodon, Argenteohyla, Corythomantis, Osteocephalus,Osteopilus, Tepuihyla, and Trachycephalus. The fourth major clade is composed ofmost of the Middle American/Holarctic species groups of Hyla and the genera Acris, Anotheca,Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca, and Triprion. Anew monophyletic taxonomy mirroring these results is presented where Hylinae is dividedinto four tribes. Of the species currently in Hyla, 297 of the 353 species are placed in 15genera; of these, 4 are currently recognized, 4 are resurrected names, and 7 are new. Hyla isrestricted to H. femoralis and the H. arborea, H. cinerea, H. eximia, and H. versicolor groups,whose contents are redefined. Phrynohyas is placed in the synonymy of Trachycephalus, andPternohyla is placed in the synonymy of Smilisca. The genus Dendropsophus is resurrectedto include all former species of Hyla known or suspected to have 30 chromosomes. Exerodontais resurrected to include the former Hyla sumichrasti group and some members of the formerH. miotympanum group. Hyloscirtus is resurrected for the former Hyla armata, H. bogotensis,and H. larinopygion groups. Hypsiboas is resurrected to include several species groupsmanyof them redefined hereof Gladiator Frogs. The former Hyla albofrenata and H. albosignatacomplexes of the H. albomarginata group are included in Aplastodiscus.New generic names are erected for (1) Agalychnis calcarifer and A. craspedopus; (2) Osteocephaluslangsdorffii; the (3) Hyla aromatica, (4) H. bromeliacia, (5) H. godmani, (6) H.mixomaculata, (7) H. taeniopus, (8) and H. tuberculosa groups; (9) the clade composed ofthe H. pictipes and H. pseudopuma groups; and (10) a clade composed of the H. circumdata,H. claresignata, H. martinsi, and H. pseudopseudis groups.