Project 4637: M. O. Pereyra, B. L. Blotto, D. Baldo, J. C. Chaparro, S. R. Ron, A. J. Elias-Costa, P. P. Iglesias, P. J. Venegas, M. T. C. Thomé, J. J. Ospina-Sarria, N. M. Maciel, M. Rada, F. Kolenc, C. Borteiro, M. Rivera-Correa, F. J. Rojas-Runjaic, J. Moravec, I. De La Riva, W. C. Wheeler, S. Castroviejo-Fisher, T. Grant, C. F. Haddad, J. Faivovich. 2021. Evolution in the Genus Rhinella: A Total Evidence Phylogenetic Analysis of Neotropical True Toads (Anura: Bufonidae). Bulletin of the American Museum of Natural History. 447 (1):1-155.
Abstract
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.Read the article »
Article DOI: 10.1206/0003-0090.447.1.1
Project DOI: 10.7934/P4637, http://dx.doi.org/10.7934/P4637
This project contains | Matrices |
---|---|
Download Project SDD File | Total scored cells: 5648 Total media associated with cells: 0 Total labels associated with cell media: 0 |
Characters | |
Total characters: 90 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 226 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:1/89 |
Currently Viewing:
MorphoBank Project 4637
MorphoBank Project 4637
- Creation Date:
12 April 2023 - Publication Date:
12 April 2023 - Project views: 4230
- Matrix downloads: 1
Authors' Institutions
- Museo de Historia Natural La Salle, Caracas
- Universidad de Antioquia
- Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)
- Universidad de Buenos Aires
- Museo Argentino de Ciencias Naturales Bernardino Rivadavia
- American Museum of Natural History
- Pontificia Universidade Catolica do Rio Grande do Sul (PUCRS)
- Universidade de Sao Paulo (USP, University of Sao Paulo)
- Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP, São Paulo State University)
- National Museum in Prague
- Universidad Nacional San Antonio Abad del Cusco
- Museo Nacional de Ciencias Naturales de Madrid (CSIC)
- Pontificia Universidad Católica del Ecuador
- Universidade Federal de Goiás
- Centro de Ornitología y Biodiversidad (CORBIDI), Peru
- Museo Nacional de Historia Natural, Montevideo
- Fundacion Calima
- Universidad Nacional de Misiones
- Museo de Biodiversidad del Perú
Members
member name | taxa | specimens | media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
MorphoBank Curator Project Administrator | 106 | 6 | 6 | 90 | 0 | 0 | 5648 (5572, 0, 76) | 0 | 0 | 0 | ||||
Martin Pereyra Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M29246' (106 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] Rhinella arunco KU217369 Last Modified in 04/12/23 | 8 | 81 | 80 | 0 | 2 | 0 | 0 | 2 |
[2] Rhinella atacamensis KU217352 Last Modified in 04/12/23 | 25 | 64 | 63 | 0 | 2 | 0 | 0 | 2 |
[3] Rhinella rubropunctata MACN52275 Last Modified in 04/12/23 | 14 | 74 | 74 | 0 | 2 | 0 | 0 | 2 |
[4] Rhinella casconi CFBH22863 Last Modified in 04/12/23 | 54 | 37 | 36 | 0 | 0 | 0 | 0 | 2 |
[5] Rhinella crucifer CFBH24630 Last Modified in 04/12/23 | 12 | 79 | 78 | 0 | 0 | 0 | 0 | 2 |
[6] Rhinella henseli CFBH20117 Last Modified in 04/12/23 | 48 | 43 | 42 | 0 | 0 | 0 | 0 | 2 |
[7] Rhinella inopina CHUNB51110 Last Modified in 04/12/23 | 69 | 21 | 21 | 0 | 0 | 0 | 0 | 2 |
[8] Rhinella ornata CFBH18815 Last Modified in 04/12/23 | 16 | 76 | 74 | 0 | 0 | 0 | 0 | 2 |
[9] Rhinella azarai LGE8710 Last Modified in 04/12/23 | 25 | 67 | 65 | 0 | 0 | 0 | 0 | 2 |
[10] Rhinella bergi MACN46555 Last Modified in 04/12/23 | 22 | 69 | 68 | 0 | 0 | 0 | 0 | 2 |
[11] Rhinella bernardoi UNSJ5046 NuG Last Modified in 04/12/23 | 49 | 43 | 41 | 0 | 0 | 0 | 0 | 2 |
[12] Rhinella beebei CBA5732 Last Modified in 04/12/23 | 35 | 56 | 55 | 0 | 0 | 0 | 0 | 2 |
[13] Rhinella dorbignyi MACN43695 Last Modified in 04/12/23 | 49 | 43 | 41 | 0 | 0 | 0 | 0 | 2 |
[14] Rhinella dorbignyi LGE8718 Last Modified in 04/12/23 | 13 | 79 | 77 | 0 | 0 | 0 | 0 | 2 |
[15] Rhinella granulosa CFBH18706 Last Modified in 04/12/23 | 49 | 41 | 41 | 0 | 0 | 0 | 0 | 2 |
[16] Rhinella humboldti AJC3533 Last Modified in 04/12/23 | 34 | 56 | 56 | 0 | 0 | 0 | 0 | 2 |
[17] Rhinella major MNCNADN6232 Last Modified in 04/12/23 | 20 | 71 | 70 | 0 | 0 | 0 | 0 | 2 |
[18] Rhinella merianae CFBH16641 Last Modified in 04/12/23 | 33 | 57 | 57 | 0 | 0 | 0 | 0 | 2 |
[19] Rhinella mirandaribeiroi CFBH10254 Last Modified in 04/12/23 | 22 | 70 | 68 | 0 | 0 | 0 | 0 | 2 |
[20] Rhinella pygmaea CFBH2894 Last Modified in 04/12/23 | 20 | 72 | 70 | 0 | 0 | 0 | 0 | 2 |
[21] Rhinella acrolopha MAR1426 Last Modified in 04/12/23 | 36 | 54 | 51 | 0 | 3 | 0 | 0 | 2 |
[22] Rhinella arborescandens MUBI14076 Last Modified in 04/12/23 | 64 | 30 | 26 | 0 | 0 | 0 | 0 | 2 |
[23] Rhinella chavin MTD43789 Last Modified in 04/12/23 | 40 | 48 | 48 | 0 | 2 | 0 | 0 | 2 |
[24] Rhinella festae KU217501 Last Modified in 04/12/23 | 38 | 50 | 49 | 0 | 3 | 0 | 0 | 2 |
[25] Rhinella lilyrodriguezae MUSM32205 Last Modified in 04/12/23 | 67 | 27 | 23 | 0 | 0 | 0 | 0 | 2 |
[26] Rhinella lindae MAR3432 Last Modified in 04/12/23 | 58 | 37 | 32 | 0 | 0 | 0 | 0 | 2 |
[27] Rhinella macrorhina MHUA8319 Last Modified in 04/12/23 | 41 | 50 | 48 | 0 | 1 | 0 | 0 | 2 |
[28] Rhinella manu MUBI11372 Last Modified in 04/12/23 | 41 | 48 | 47 | 0 | 2 | 0 | 0 | 2 |
[29] Rhinella cf multiverrucosa MUBI11455 Last Modified in 04/12/23 | 39 | 51 | 49 | 0 | 2 | 0 | 0 | 2 |
[30] Rhinella nesiotes CORBIDI13953 Last Modified in 04/12/23 | 69 | 25 | 21 | 0 | 0 | 0 | 0 | 2 |
[31] Rhinella cf nicefori MHUA4793 Last Modified in 04/12/23 | 60 | 31 | 29 | 0 | 1 | 0 | 0 | 2 |
[32] Rhinella paraguas TG1480 Last Modified in 04/12/23 | 46 | 46 | 43 | 0 | 1 | 0 | 0 | 2 |
[33] Rhinella ruizi AML40 Last Modified in 04/12/23 | 62 | 27 | 25 | 0 | 3 | 0 | 0 | 2 |
[34] Rhinella tacana MNK7187 Last Modified in 04/12/23 | 64 | 30 | 26 | 0 | 0 | 0 | 0 | 2 |
[35] Rhinella tenrec MAR3584 Last Modified in 04/12/23 | 66 | 23 | 21 | 0 | 3 | 0 | 0 | 2 |
[36] Rhinella yanachaga MUBI7119 Last Modified in 04/12/23 | 45 | 47 | 45 | 0 | 0 | 0 | 0 | 2 |
[37] Rhinella acutirostris MTR36593 Last Modified in 04/12/23 | 66 | 26 | 23 | 0 | 1 | 0 | 0 | 2 |
[38] Rhinella alata QCAZ11597 Last Modified in 04/12/23 | 60 | 34 | 30 | 0 | 0 | 0 | 0 | 2 |
[39] Rhinella castaneotica LSUMZ17429 Last Modified in 04/12/23 | 31 | 59 | 58 | 0 | 1 | 0 | 0 | 2 |
[40] Rhinella dapsilis QCAZ43967 Last Modified in 04/12/23 | 34 | 56 | 56 | 0 | 0 | 0 | 0 | 2 |
[41] Rhinella dapsilis CFBH11400 Last Modified in 04/12/23 | 66 | 25 | 24 | 0 | 0 | 0 | 0 | 2 |
[42] Rhinella hoogmoedi CFBH13286 Last Modified in 04/12/23 | 37 | 54 | 52 | 0 | 1 | 0 | 0 | 2 |
[43] Rhinella lescurei MC5 Last Modified in 04/12/23 | 68 | 26 | 22 | 0 | 0 | 0 | 0 | 2 |
[44] Rhinella magnussoni APL20530 Last Modified in 04/12/23 | 51 | 42 | 37 | 0 | 2 | 0 | 0 | 2 |
[45] Rhinella sp 10 QCAZ42269 Last Modified in 04/12/23 | 51 | 40 | 39 | 0 | 0 | 0 | 0 | 2 |
[46] Rhinella sp 14 KU215146 Last Modified in 04/12/23 | 46 | 46 | 44 | 0 | 0 | 0 | 0 | 2 |
[47] Rhinella margaritifera MC156 Last Modified in 04/12/23 | 68 | 26 | 22 | 0 | 0 | 0 | 0 | 2 |
[48] Rhinella ocellata ZUFG8519 Last Modified in 04/12/23 | 27 | 65 | 63 | 0 | 0 | 0 | 0 | 2 |
[49] Rhinella scitula UFMT1876 Last Modified in 04/12/23 | 61 | 34 | 29 | 0 | 0 | 0 | 0 | 2 |
[50] Rhinella proboscidea CTGAUFAM5602 Last Modified in 04/12/23 | 56 | 35 | 32 | 0 | 2 | 0 | 0 | 2 |
[51] Rhinella scitula IIBP849 Last Modified in 04/12/23 | 51 | 44 | 39 | 0 | 0 | 0 | 0 | 2 |
[52] Rhinella sclerocephala MHNLS7495 Last Modified in 04/12/23 | 64 | 28 | 26 | 0 | 0 | 0 | 0 | 2 |
[53] Rhinella stanlaii MNCN ADN6274 Last Modified in 04/12/23 | 62 | 31 | 26 | 0 | 2 | 0 | 0 | 2 |
[54] Rhinella sp 5 QCAZ53142 Last Modified in 04/12/23 | 69 | 25 | 21 | 0 | 0 | 0 | 0 | 2 |
[55] Rhinella iserni MUSM31096 Last Modified in 04/12/23 | 50 | 40 | 38 | 0 | 2 | 0 | 0 | 2 |
[56] Rhinella achavali MNHN Uy9301 Last Modified in 04/12/23 | 36 | 54 | 54 | 0 | 0 | 0 | 0 | 2 |
[57] Rhinella arenarum AR305 Last Modified in 04/12/23 | 14 | 78 | 76 | 0 | 0 | 0 | 0 | 2 |
[58] Rhinella cerradensis CHUNB38671 Last Modified in 04/12/23 | 24 | 66 | 66 | 0 | 0 | 0 | 0 | 2 |
[59] Rhinella diptycha KU289057 Last Modified in 04/12/23 | 21 | 72 | 69 | 0 | 0 | 0 | 0 | 2 |
[60] Rhinella horribilis MAR2057 NuG Last Modified in 04/12/23 | 30 | 63 | 60 | 0 | 0 | 0 | 0 | 2 |
[61] Rhinella icterica CFBH27410 Last Modified in 04/12/23 | 25 | 68 | 65 | 0 | 0 | 0 | 0 | 2 |
[62] Rhinella diptycha CFBH19523 Last Modified in 04/12/23 | 18 | 73 | 72 | 0 | 0 | 0 | 0 | 2 |
[63] Rhinella marina VUB1965 Last Modified in 04/12/23 | 24 | 69 | 66 | 0 | 0 | 0 | 0 | 2 |
[64] Rhinella poeppigii MUBI6863 Last Modified in 04/12/23 | 38 | 55 | 52 | 0 | 0 | 0 | 0 | 2 |
[65] Rhinella rubescens CFBH5836 Last Modified in 04/12/23 | 21 | 69 | 69 | 0 | 0 | 0 | 0 | 2 |
[66] Rhinella veredas CHUNB44609 Last Modified in 04/12/23 | 46 | 46 | 44 | 0 | 0 | 0 | 0 | 2 |
[67] Rhinella achalensis MACN52406 Last Modified in 04/12/23 | 31 | 59 | 57 | 0 | 2 | 0 | 0 | 2 |
[68] Rhinella cf amabilis QCAZ68471 Last Modified in 04/12/23 | 39 | 50 | 50 | 0 | 1 | 0 | 0 | 2 |
[69] Rhinella spinulosa KU214792 Last Modified in 04/12/23 | 32 | 63 | 58 | 0 | 0 | 0 | 0 | 2 |
[70] Rhinella gallardoi LGE4546 Last Modified in 04/12/23 | 41 | 48 | 47 | 0 | 2 | 0 | 0 | 2 |
[71] Rhinella limensis nv Last Modified in 04/12/23 | 32 | 58 | 55 | 0 | 3 | 0 | 0 | 2 |
[72] Rhinella papillosa MACN49782 Last Modified in 04/12/23 | 52 | 40 | 36 | 0 | 2 | 0 | 0 | 2 |
[73] Rhinella altiperuviana MACN49701 Last Modified in 04/12/23 | 19 | 74 | 71 | 0 | 0 | 0 | 0 | 2 |
[74] Rhinella spinulosa nv Last Modified in 04/12/23 | 32 | 58 | 55 | 0 | 3 | 0 | 0 | 2 |
[75] Rhinella trifolium nv Last Modified in 04/12/23 | 63 | 27 | 27 | 0 | 0 | 0 | 0 | 2 |
[76] Rhinella vellardi KU211765 Last Modified in 04/12/23 | 39 | 48 | 48 | 0 | 3 | 0 | 0 | 2 |
[77] Rhinella sternosignata nv Last Modified in 04/12/23 | 40 | 51 | 50 | 0 | 0 | 0 | 0 | 2 |
[78] Rhinella quechua MNK5302 Last Modified in 04/12/23 | 68 | 23 | 20 | 0 | 2 | 0 | 0 | 2 |
[79] Rhinella inca MNCN44405 Last Modified in 04/12/23 | 72 | 20 | 18 | 0 | 0 | 0 | 0 | 2 |
[80] Rhinella justinianoi MNCN ADN6065 Last Modified in 04/12/23 | 67 | 23 | 23 | 0 | 0 | 0 | 0 | 2 |
[81] Rhinella leptoscelis MUBI5989 Last Modified in 04/12/23 | 72 | 19 | 18 | 0 | 0 | 0 | 0 | 2 |
[82] Rhinella quechua MNCN ADN3927 Last Modified in 04/12/23 | 12 | 79 | 75 | 0 | 3 | 0 | 0 | 2 |
[83] Rhinella rumbolli MACN53782 Last Modified in 04/12/23 | 7 | 83 | 83 | 0 | 0 | 0 | 0 | 2 |
[84] Rhinella veraguensis USNM346048 Last Modified in 04/12/23 | 34 | 56 | 56 | 0 | 0 | 0 | 0 | 2 |
[85] Amazophrynella minuta Last Modified in 04/12/23 | 30 | 58 | 56 | 0 | 4 | 0 | 0 | 2 |
[86] Anaxyrus boreas Last Modified in 04/12/23 | 24 | 68 | 65 | 0 | 1 | 0 | 0 | 2 |
[87] Anaxyrus americanus Last Modified in 04/12/23 | 20 | 74 | 70 | 0 | 0 | 0 | 0 | 2 |
[88] Anaxyrus woodhousii Last Modified in 04/12/23 | 20 | 72 | 70 | 0 | 0 | 0 | 0 | 2 |
[89] Ansonia longidigita Last Modified in 04/12/23 | 50 | 45 | 39 | 0 | 1 | 0 | 0 | 2 |
[90] Bufo bufo Last Modified in 04/12/23 | 17 | 73 | 70 | 0 | 3 | 0 | 0 | 2 |
[91] Duttaphrynus melanostictus Last Modified in 04/12/23 | 24 | 68 | 66 | 0 | 0 | 0 | 0 | 2 |
[92] Incilius alvarius Last Modified in 04/12/23 | 28 | 64 | 61 | 0 | 1 | 0 | 0 | 2 |
[93] Incilius coniferus Last Modified in 04/12/23 | 26 | 68 | 64 | 0 | 0 | 0 | 0 | 2 |
[94] Incilius valliceps Last Modified in 04/12/23 | 23 | 69 | 67 | 0 | 0 | 0 | 0 | 2 |
[95] Melanophryniscus stelzneri Last Modified in 04/12/23 | 9 | 82 | 79 | 0 | 2 | 0 | 0 | 2 |
[96] Nannophryne variegata Last Modified in 04/12/23 | 12 | 78 | 75 | 0 | 3 | 0 | 0 | 2 |
[97] Peltophryne lemur Last Modified in 04/12/23 | 35 | 58 | 55 | 0 | 0 | 0 | 0 | 2 |
[98] Peltophryne empusa Last Modified in 04/12/23 | 37 | 55 | 53 | 0 | 0 | 0 | 0 | 2 |
[99] Phrynoidis juxtaspera Last Modified in 04/12/23 | 26 | 68 | 64 | 0 | 0 | 0 | 0 | 2 |
[100] Rentapia hosii Last Modified in 04/12/23 | 24 | 69 | 66 | 0 | 0 | 0 | 0 | 2 |
[101] Rhaebo guttatus Last Modified in 04/12/23 | 17 | 75 | 73 | 0 | 0 | 0 | 0 | 2 |
[102] Rhaebo ceratophrys QCAZ40240 Last Modified in 04/12/23 | 57 | 34 | 33 | 0 | 0 | 0 | 0 | 2 |
[103] Schismaderma carens Last Modified in 04/12/23 | 22 | 68 | 66 | 0 | 2 | 0 | 0 | 2 |
[104] Sclerophrys mauritanica Last Modified in 04/12/23 | 44 | 49 | 46 | 0 | 0 | 0 | 0 | 2 |
[105] Sclerophrys regularis Last Modified in 04/12/23 | 12 | 84 | 78 | 0 | 0 | 0 | 0 | 2 |
[106] Vandijkophrynus robinsoni Last Modified in 04/12/23 | 52 | 42 | 38 | 0 | 0 | 0 | 0 | 2 |
Project views
type | number of views | Individual items viewed (where applicable) |
Total project views | 4230 | |
Project overview | 415 | |
Matrix views | 173 | Matrix landing page (168 views); Matrix (5 views); |
Documents list | 175 | |
Specimen list | 748 | |
Bibliography | 144 | |
Taxon list | 1168 | |
Views for media list | 214 | |
Media views | 1193 | Media search (573 views); M861418 (107 views); M861419 (97 views); M861420 (112 views); M861422 (103 views); M861423 (102 views); M861421 (99 views); |
Project downloads
type | number of downloads | Individual items downloaded (where applicable) |
Total downloads from project | 6 | |
Project downloads | 5 | |
Matrix downloads | 1 | Matrix (1 download); |