A new genus of fossil frog (Anura) from lower Cretaceous deposits in South America

The frog clade Neobatrachia or “advanced frogs” represents more than 95% of the diversity within Anura. The neobatrachian fossil record is incomplete due to the small size and fragile nature of their bones but provides some clues on the time and rate of the current diversification of modern-day amphibians. The Crato Formation (Aptian) of northeastern Brazil is known for the presence of, at least, five species of neobatrachian fossils. Herein, we describe a well-preserved fossil that represents a new neobatrachian taxon from the Crato Formation (Cretaceous) in the Araripe Basin, northeastern Brazil, and perform phylogenetic analysis to assess its higher-level relationships. The new specimen is an almost complete and articulated skeleton with soft tissue preservation. Phylogenetic analysis recovered the specimen nested in the clade Hyloidea and our results highlight the remarkable value of the Crato Formation as a Laggerstätte.

Key words
Amphibia; fossil anuran; Cretaceous; Primaevorana cratensis

Authorship

PEDRO HENRIQUE A.G. MOURA

Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP), Laboratory of Herpetology, Institute of Biosciences, Campus do Litoral Paulista, Pça. Infante D. Henrique, s/n, Parque Bitaru, 11330-900 São Vicente, SP, BrazilUniversidade Estadual Paulista Júlio de Mesquita Filho (UNESP)BrazilSão Vicente, SP, BrazilUniversidade Estadual Paulista Júlio de Mesquita Filho (UNESP), Laboratory of Herpetology, Institute of Biosciences, Campus do Litoral Paulista, Pça. Infante D. Henrique, s/n, Parque Bitaru, 11330-900 São Vicente, SP, Brazil

http://orcid.org/0000-0003-3391-6149

FABIANA R. COSTA

Universidade Federal do ABC (UFABC), Laboratory of Vertebrate Paleontology and Animal Behavior (LAPC), Center of Natural and Human Sciences, Campus São Bernardo do Campo, Alameda da Universidade, s/n, Anchieta, 09606-045 São Bernardo do Campo, SP, BrazilUniversidade Federal do ABC (UFABC)BrazilSão Bernardo do Campo, SP, BrazilUniversidade Federal do ABC (UFABC), Laboratory of Vertebrate Paleontology and Animal Behavior (LAPC), Center of Natural and Human Sciences, Campus São Bernardo do Campo, Alameda da Universidade, s/n, Anchieta, 09606-045 São Bernardo do Campo, SP, Brazil

http://orcid.org/0000-0003-3596-0143

LUIZ E. ANELLI

Universidade de São Paulo (USP), Department of Sedimentary and Environmental Geology, Geosciences Institute, Rua do Lago, 562, Butantã, 05508-900 São Paulo, SP, BrazilUniversidade de São Paulo (USP)BrazilSão Paulo, SP, BrazilUniversidade de São Paulo (USP), Department of Sedimentary and Environmental Geology, Geosciences Institute, Rua do Lago, 562, Butantã, 05508-900 São Paulo, SP, Brazil

http://orcid.org/0000-0002-5228-9833

IVAN NUNES

http://orcid.org/0000-0001-7985-2836

Correspondence to: Fabiana Rodrigues Costa E-mail: fabiana.costa@ufabc.edu.br

AUTHOR CONTRIBUTIONS

Costa FR, Anelli LE, and Nunes I conceived the study. Moura PHAG, Costa FR, and Nunes I delimited the methods and analyses. Moura PHAG and Nunes I carried out the analyses and interpreted the results. All authors worked on the manuscript and contributed to the final version and revisions.

SCIMAGO INSTITUTIONS RANKINGS

Figures

Figures (6)

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Figure 1
Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Photograph by Luiz E. Anelli; b) Illustration of the fossil. Abbreviations: pmp: posteromedial processes of the hyoid; pho: parahyoid ossification; ps: parasphenoid; 3+4+5: distal carpals 3+4+5; ps: palmar sesamoid; y+2: element y + distal carpal 2; p: proximal element of the prepollex; 2+3: distal tarsals 2+3; pha: prehallux.

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Figure 2
Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Cranium; b) Pectoral girdle; c) Anterior portion of the cranium; d) Trunk. Photographs by Ivan Nunes. Abbreviations: pmp: posteromedial processes of the hyoid; pho: parahyoid ossification; ps: parasphenoid.

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Figure 3
Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Right forelimb; b) Left forelimb. Photographs by Ivan Nunes. Abbreviations: 3+4+5: distal carpals 3+4+5; ps: palmar sesamoid; y+2: element y + distal carpal 2; p: proximal element of the prepollex.

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Figure 4
Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Pelvic girdle; b) Detail view of pelvic girdle; c) Right hindlimb; d) Left hindlimb. Photographs by Ivan Nunes.

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Figure 5
Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Right hindlimb; b) Left Hindlimb; c) Right pes; d) Left pes. Photographs by Ivan Nunes. Abbreviations: 2+3: distal tarsals 2+3; pha: prehallux.

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Figure 6
Phylogenetic trees obtained from the maximum parsimony analyses using the heuristic search of TNT v. 1.1, under implied weights. a) Unconstrained analysis with k = 7; b) Constrained analysis with k = 15. The fossil taxa are indicated with an asterisk. The new species is highlighted in bold. Numbers above nodes are relative Goodman-Bremer support values; numbers below nodes are Jackknife values. Only Jackknife values above 50% are displayed.

Figure 1 Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Photograph by Luiz E. Anelli; b) Illustration of the fossil. Abbreviations: pmp: posteromedial processes of the hyoid; pho: parahyoid ossification; ps: parasphenoid; 3+4+5: distal carpals 3+4+5; ps: palmar sesamoid; y+2: element y + distal carpal 2; p: proximal element of the prepollex; 2+3: distal tarsals 2+3; pha: prehallux.

Figure 2 Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Cranium; b) Pectoral girdle; c) Anterior portion of the cranium; d) Trunk. Photographs by Ivan Nunes. Abbreviations: pmp: posteromedial processes of the hyoid; pho: parahyoid ossification; ps: parasphenoid.

Figure 3 Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Right forelimb; b) Left forelimb. Photographs by Ivan Nunes. Abbreviations: 3+4+5: distal carpals 3+4+5; ps: palmar sesamoid; y+2: element y + distal carpal 2; p: proximal element of the prepollex.

Figure 4 Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Pelvic girdle; b) Detail view of pelvic girdle; c) Right hindlimb; d) Left hindlimb. Photographs by Ivan Nunes.

Figure 5 Primaevorana cratensis, holotype (GP/2E-9497) from Crato Formation, Lower Cretaceous (Aptian), region of Nova Olinda, Araripe Basin, Ceará state, Northeastern Brazil. a) Right hindlimb; b) Left Hindlimb; c) Right pes; d) Left pes. Photographs by Ivan Nunes. Abbreviations: 2+3: distal tarsals 2+3; pha: prehallux.

Figure 6 Phylogenetic trees obtained from the maximum parsimony analyses using the heuristic search of TNT v. 1.1, under implied weights. a) Unconstrained analysis with k = 7; b) Constrained analysis with k = 15. The fossil taxa are indicated with an asterisk. The new species is highlighted in bold. Numbers above nodes are relative Goodman-Bremer support values; numbers below nodes are Jackknife values. Only Jackknife values above 50% are displayed.

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A new genus of fossil frog (Anura) from lower Cretaceous deposits in South America

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[1] Correspondence to: Fabiana Rodrigues Costa E-mail: fabiana.costa@ufabc.edu.br