Evolution in the Genus Rhinella: a total evidence phylogenetic analysis of neotropical true toads (Anura: Bufonidae) (2021)
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2021
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True toads of the genus Rhinella are among the most common and diverse group of Neotropical
anurans. These toads are widely distributed throughout South America, inhabiting a great diversity
of environments and ecoregions. Currently, however, the genus is defined solely on the basis of
molecular characters, and it lacks a proper diagnosis. Although some phenetic species groups have
traditionally been recognized within Rhinella, the monophyly of some of them have been rejected
in previous phylogenetic analyses, and many species remain unassigned to these poorly defined
groups. Additionally, the identity and taxonomy of several species are problematic and hinder the
specific recognition and description of undescribed taxa. In this work, we first perform phylogenetic
analyses of separate mitochondrial and nuclear datasets to test the possible occurrence of hybridization and/or genetic introgression in the genus. The comparative analysis of both datasets revealed
unidirectional mitochondrial introgressions of an unknown parental species into R. horribilis (“ghost
introgression”) and of R. dorbignyi into R. bernardoi; therefore, the mitochondrial and nuclear datasets of these species were considered separately in subsequent analyses. We performed total-evidence
phylogenetic analyses that included revised molecular (four mitochondrial and five nuclear genes)
and phenotypic (90 characters) datasets for 83 nominal species of Rhinella, plus several undescribed
and problematic species and multiple outgroups. Results demonstrate that Rhinella was nonmonophyletic due to the position of R. ceratophrys, which was recovered as the sister taxon of Rhaebo
nasicus with strong support. Among our outgroups, the strongly supported Anaxyrus + Incilius is
the sister clade of all other species of Rhinella. Once R. ceratophrys is excluded, the genus Rhinella
is monophyletic, well supported, and composed of two major clades. One of these is moderately
supported and includes species of the former R. spinulosa Group (including R. gallardoi); the monophyletic R. granulosa, R. crucifer, and R. marina Groups; and a clade composed of the mitochondrial
sequences of R. horribilis. The other major clade is strongly supported and composed of all the species from the non-monophyletic R. veraguensis and R. margaritifera Groups, the former R. acrolopha
Group, and R. sternosignata. Consistent with these results, we define eight species groups of Rhinella
that are mostly diagnosed by phenotypic synapomorphies in addition to a combination of morphological character states. Rhinella sternosignata is the only species that remains unassigned to any
group. We also synonymize nine species, treat three former subspecies as full species, and suggest
that 15 lineages represent putative undescribed species. Lastly, we discuss the apparently frequent
occurrence of hybridization, deep mitochondrial divergence, and “ghost introgression”; the incomplete phenotypic evidence (including putative character systems that could be used for future phylogenetic analyses); and the validity of the known fossil record of Rhinella as a source of calibration
points for divergence dating analyses.
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PEREYRA, Martín O. et al. Evolution in the Genus Rhinella: a total evidence phylogenetic analysis of neotropical true toads (Anura: Bufonidae) (2021). Bulletin of the American Museum of Natural History, Washington, v. 447, n. 1, p. 1-155, 2021. DOI: 10.1206/0003-0090.447.1.1. Disponível em: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-447/issue-1#Articles. Acesso em: 20 jul. 2023.