Frogs of the Hyla arntata and Hyla pulchella Groups in the Andes of South America, with Definitions and Analyses of Phylogenetic Relationships of Andean Groups of Hyla

Abstract

Andean hylid frogs of the genus Hyla are reviewed. Eight groups of Hyla recognized in the Andes are defined; the species in two of these groups are treated in detail. The large H. armata, characterized by hypertrophied forelimbs and clumps of nuptial spines on the prepoUex and humerus in males, and a stream-adapted tadpole with a LTRF up to 15/17, is the sole recognized species in the H. armata group. Accounts of this species and six species (one new) in the H. piilchella group include diagnoses, descriptions of adults, larvae, advertisement calls, and information on ecology, and distribution. Hyla ocapia Andersson, 1938, is placed in the synonymy of H. albonigra Nieden, 1923; H. callipkura Boulenger 1902, is placed in the synonymy of H. balzani Boulenger 1898, and H. pulchella andina Muller, 1924, is recognized as a distinct species. The H. pulchella group is characterized by hypertrophied forelimbs and projecting prepollical spines in adult males, and generalized, stream-adapted tadpoles. These characters are shared by frogs in the H. circumdata group in southeastern Brazil, and the two groups are combined herein into the H. pulchella group. The presence of sharp, projecting prepollical spines is a character shared with several groups of Hyla in the neotropical lowlands. Phylogenetic analysis reveals that the H. columbiana and H. labialis groups are sister groups; this clade is the sister of a clade containing several groups of Hyla having a diploid number of 30 chromosomes and inhabiting Andes (H. garagoensis Group) and the neotropical lowlands. Frogs in the H. bogotensis group are characterized by the presence of a mental gland in males; this derived character is shared with H. granosa in the Amazon Basin and the Guiana Region. No unique synapomorphies exist for the frogs assigned to the H. larinopygion group. Too few data are available for Hyla chlorostea to associate it with any lineages of Hyla.