Dryophytes Fitzinger, 1843

Duellman, William E., Marion, Angela B. & Hedges, Blair, 2016, Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae), Zootaxa 4104 (1), pp. 1-109 : 23

Dryophytes Fitzinger, 1843

Dryophytes Fitzinger, 1843:31 . Type species: Hyla versicolor LeConte, 1825 , by original designation.

Definition. Moderate-sized, arboreal, primarily green frogs with expanded terminal discs on the digits ( Figs. 8 C and D); no definitive morphological features are known to separate Dryophytes from Hyla .

Content. Nineteen species: Dryophytes andersonii (Baird) , arboricola* (Taylor), arenicolor (Cope) , avivocus (Viosca), bocourti* (Mocquard), chrysoscelis (Cope) , cinereus (Schneider), euphorbiaceus (Günther), eximius (Baird), femoralis (Daudin) , gratiosus (LeConte), immaculatus Boettger, japonicus (Günther) , plicatus (Brocchi), squirellus (Daudin), suweonensis (Kuramoto) , versicolor (LeConte) , walkeri (Stuart) , and wrightorum (Taylor) ; all new combinations.

Distribution. North America east of the Sierra Nevada southward from extreme southern Canada to the Gulf of Mexico and on the Mexican Plateau southward to Oaxaca, plus the highlands of Chiapas, Mexico, and adjacent Guatemala. Nearctic Region in far eastern Russia, Korean Peninsula, Japan, eastern China, and Ryukyu Island

Etymology. The generic name is derived from the Greek dryos meaning tree and the Greek phytes meaning plant. This rather redundant epithet presumably refers to the arboreal habits of these frogs. The gender is masculine.

Remarks. The inclusion of Asian and North American taxa in the same genus is like the biogeography of Rana , a genus with 41 species in Eurasia and seven species in western North America (Frost 2015).

Sixteen species of Dryophytes occur is eastern North America, and three species are found in eastern temperate Asia. We purposefully excluded sequences in Genbank ( FJ226937 , FJ226830 ) identified as “ Hyla heinzsteinitzi ” because they were labeled incorrectly. They are Dryophytes japonica , introduced to Israel, as was determined by the authors of the study that generated the sequences ( Stöck et al. 2008).

In the trees produced by Faivovich et al. (2005) and Hua et al. (2009), a sample identified as “ Hyla ” walkeri , a species restricted to the highlands of western Guatemala and adjacent Mexico, is the sister species of “ Hyla ” immaculata , a species occurring in eastern China. Hua et al. (2009:256) examined the voucher specimen of “ Hyla ” walkeri used by Faivovich et al. (2005); they noted that this specimen (AMNH-A 168406) came from the pet trade, has no locality data, and closely resembled specimens of “ Hyla ” immaculata from China. The true “ Hyla ” walkeri sample included in Hua et al. (2009) falls out in a clade of five Mexican species, all members of the Hyla eximia Group recognized by Duellman (2001). In our analysis, we included only those GenBank sequences positively identified as “ Hyla ” walkeri , from Hua et al. (2009) and Lemmon et al. (2007). (Pyron and Wiens [2011] included sequences from both the pet-trade “ walkeri *” sample from Faivovich et al. [2005] and the true walkeri sample from Hua et al. [2009] as a single chimeric taxon.) Thus in our analysis ( Fig. 4 ), Dryophytes walkeri is a member of the Hyla eximia Group recognized by Duellman (2001).

Figures

FIGURE 4. Maximum likelihood phylogeny of arboranan frogs based on DNA sequence data from 19 genes (16,128 aligned sites). The tree is rooted with Ceuthomantis smaragdinus, Dendrobates auratus, Haddadus binotatus, and Rhinoderma darwinii (not shown). Bootstrap support values are indicated at nodes.

FIGURE 8. A. Hyla arborea, Tubingen, Baden Württenburg, Germany. B. Hyla sarda, KU 207373, 2.4 km ESE Musei (near Domusnovas), Cagliari, Sardinia, Italy. C. Dryophytes versicolor, near Greenville, Darke County, Ohio, USA. D. Dryophytes gratiosus, KU 109911, Woodland, East Feliciana Parish, Louisiana, USA. All by W. E. Duellman.