SYSTEMATIC REVIEW OF THE FROG FAMILY HYLIDAE, WITH SPECIAL REFERENCE TO HYLINAE: PHYLOGENETIC ANALYSIS AND TAXONOMIC REVISION

By J. Faivovich, Célio Fernando Baptista Haddad, P. Garcia, D. Frost, J. Campbell, Ward C. Wheeler. 2005

999 citations recorded by the source corpus.

Abstract Hylidae is a large family of American, Australopapuan, and temperate Eurasian treefrogs of approximately 870 known species, divided among four subfamilies. Although some groups of Hylidae have been addressed phylogenetically, a comprehensive phylogenetic analysis has never been presented. The first goal of this paper is to review the current state of hylid systematics. We focus on the very large subfamily Hylinae (590 species), evaluate the monophyly of named taxa, and examine the evidential basis of the existing taxonomy. The second objective is to perform a phylogenetic analysis using mostly DNA sequence data in order to (1) test the monophyly of the Hylidae; (2) determine its constituent taxa, with special attention to the genera and species groups which form the subfamily Hylinae, and c) propose a new, monophyletic taxonomy consistent with the hypothesized relationships. We present a phylogenetic analysis of hylid frogs based on 276 terminals, including 228 hylids 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 the species-rich genus Hyla. The included taxa allowed us to test the monophyly of 24 of the 35 nonmonotypic genera and 25 species groups of Hyla. The phylogenetic analysis includes approximately 5100 base pairs from four mitochondrial (12S, tRNA valine, 16S, and cytochrome b) and five nuclear genes (rhodopsin, tyrosinase, RAG-1, seventh in absentia, and 28S), and a small data set from foot musculature. Concurring with previous studies, the present analysis indicates that Hemiphractinae are not related to the other three hylid subfamilies. It is therefore removed from the family and tentatively considered a subfamily of the paraphyletic Leptodactylidae. Hylidae is now restricted to Hylinae, Pelodryadinae, and Phyllomedusinae. Our results support a sister-group relationship between Pelodryadinae and Phyllomedusinae, which together form the sister taxon of Hylinae. 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 also demonstrably nonmonophyletic. Hylinae is composed of four major clades. The first of these includes the Andean stream-breeding Hyla, Aplastodiscus, all Gladiator Frogs, and a Tepuian clade. The second clade is composed of the 30-chromosome Hyla, Lysapsus, Pseudis, Scarthyla, Scinax (including the H. uruguaya group), Sphaenorhynchus, and Xenohyla. The third major 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 of most of the Middle American/Holarctic species groups of Hyla and the genera Acris, Anotheca, Duellmanohyla, Plectrohyla, Pseudacris, Ptychohyla, Pternohyla, Smilisca, and Triprion. A new monophyletic taxonomy mirroring these results is presented where Hylinae is divided into four tribes. Of the species currently in “Hyla”, 297 of the 353 species are placed in 15 genera; of these, 4 are currently recognized, 4 are resurrected names, and 7 are new. Hyla is restricted 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, and Pternohyla is placed in the synonymy of Smilisca. The genus Dendropsophus is resurrected to include all former species of Hyla known or suspected to have 30 chromosomes. Exerodonta is resurrected to include the former Hyla sumichrasti group and some members of the former H. miotympanum group. Hyloscirtus is resurrected for the former Hyla armata, H. bogotensis, and H. larinopygion groups. Hypsiboas is resurrected to include several species groups—many of them redefined here—of Gladiator Frogs. The former Hyla albofrenata and H. albosignata complexes of the H. albomarginata group are included in Aplastodiscus. New generic names are erected for (1) Agalychnis calcarifer and A. craspedopus; (2) Osteocephalus langsdorffii; 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 of the H. pictipes and H. pseudopuma groups; and (10) a clade composed of the H. circumdata, H. claresignata, H. martinsi, and H. pseudopseudis groups.

Subfields: Genetics, Global and Planetary Change, Molecular Biology

View Paper: https://www.semanticscholar.org/paper/57cd97772d7c090cba1ee1c3f1c66c231c5d9ad5

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Citations in the corpus

Systematics of the Neotropical Genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-molecular Evidence, and Species Accounts (2014, background): “In the 21st Century, anuran systematics and taxonomy has progressed rapidly based on both molecular and detailed morphological studies (Haas, 2003; Faivovich et al., 2005; Frost et al., 2006; Grant et al., 2006; Van der Meijen et al., 2007; Pyron and Wiens, 2011; de Sá et al., 2012).”
Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs (2007, background): “Flores (1987) described the parietal peritoneum of ametarsia as clear; however, it is also covered by iridophores for at least A of its extent (our state PI) as in oyampiensis (pers.”
A new frog family (Anura: Terrarana) from South America and an expanded direct-developing clade revealed by molecular phylogeny (2009, positive): “Species composition of the chimeric sequences was guided by the results of previous molecular phylogenetic analyses (Frost et al. 2006; Wiens et al. 2005; Faivovich et al. 2005; Grant et al. 2007; Darst and Cannatella 2004; Guayasamin et al. 2008; Roelants et al. 2007).”
Genome duplication in amphibians and fish: an extended synthesis (2011, background): “Polyploid species (and their diploid progenitors) tend to be underrepresented in molecular phylogenies (e.g. Faivovich et al., 2005; Frost et al., 2006), possibly as a result of the practical difficulties of dealing with duplicated gene sequences, but also the fact that polyploid taxa are not…”
The amphibian chytrid fungus along an altitudinal transect before the first reported declines in Costa Rica (2006, background): “The taxonomy of the species follows Savage (2002), Crawford and Smith (2005) and Faivovich et al. (2005) .”
PHYLOGENETIC SYSTEMATICS OF DART-POISON FROGS AND THEIR RELATIVES (AMPHIBIA: ATHESPHATANURA: DENDROBATIDAE) (2006, background): “Faivovich et al. (2005) were primarily interested in the relationships among hylids, but their outgroup sample was extensive; their analysis of nuclear and mitochondrial DNA placed dendrobatids as the sister group to the Hylodinae (topology not shown).”
Anurans as prey: an exploratory analysis and size relationships between predators and their prey (2007, background): “Specific names are in agreement with online databases: amphibians follow Frost (2004) complemented by Faivovich et al. (2005), Nascimento, Caramaschi & Cruz (2005) and Frost et al. (2006), reptiles follow Uetz et al. (2005), fishes follow Froese & Pauly (2004), birds follow Lepage (2005) and…”
Species delimitation, patterns of diversification and historical biogeography of the Neotropical frog genus Adenomera (Anura, Leptodactylidae) (2014, background): “Several open habitat species of other taxa are endemic to Serra do Cachimbo (plants: Zappi et al., 2011; Squamata: Rodrigues, 1987; Anura: Faivovich et al., 2005).”
A new genus and species of rodent from the Brazilian Atlantic Forest (Rodentia: Cricetidae: Sigmodontinae: Oryzomyini), with comments on oryzomyine biogeography (2011, background): “In their comprehensive systematic revisions of family Hylidae, Faivovich et al. (2004, 2005) observed that in the subgroup Hypsiboas pulchellus an ‘. . . Andean clade is nested within an Atlantic Forest/ Cerrado clade . . . a biogeographic pattern that we are not aware of in other vertebrate groups’”
Gliding and the Functional Origins of Flight: Biomechanical Novelty or Necessity? (2007, background): “…in numerous New World species of the family Hylidae, including Ecnomiohyla miliaria, Phyllomedusa callidryas, Pachymedusa dacnicolor, Agalychnis spurrelli, A. saltator, and Scinax ruber (Duellman 2001, Faivovich et al. 2005, Pauly et al. 2005, Pyburn 1970, Roberts 1994, Scott & Starrett 1974).”
Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness (COVER) (2011, background): “However, while several phylogenetic studies have suggested that these subfamilies form a clade (Faivovich et al., 2005; Frost et al., 2006; Grant et al., 2006), others”
Casting the Percomorph Net Widely: The Importance of Broad Taxonomic Sampling in the Search for the Placement of Serranid and Percid Fishes (2007, background): “Because of the large size of the data set, an extensive and thorough search strategy was necessary (Faivovich et al., 2005).”
Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of São Paulo State, Brazil (2006, methodology): “The characters are described according to Altig & McDiarmid (1999b), with coloration analysis based on preserved specimens, and the classification according Frost (2004), Faivovich et al. (2005) and Nascimento et al. (2005).”