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Type: Article
Published: 2021-12-17
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Conversion of the names Pyxicephaloidea, Pyxicephalidae and Pyxicephalinae (Anura, Ranoidea) into phylogenetic nomenclature

CR2P- Centre de recherche en Paléontologie-Paris, CNRS/MNHN/Sorbonnes Université, Bâtiment de Géologie, 43 rue Buffon, Paris, 75005, France
CR2P- Centre de recherche en Paléontologie-Paris, CNRS/MNHN/Sorbonnes Université, Bâtiment de Géologie, 43 rue Buffon, Paris, 75005, France
Phylogenetic nomenclature PhyloCode amphibians Pyxicephalidae


The recent publications of the sixth edition of the PhyloCode and of the monograph Phylonyms now allow the publication of nomenclatural acts that will establish priority under that code. This includes defining existing and newly proposed taxon names in conformity with the PhyloCode. Among amphibian taxa, very few names have been converted so far, and we take the opportunity of our recent phylogenetic analysis of neobatrachians focusing on an extinct genus of Pyxicephalidae, Thaumastosaurus, from the Eocene of Western Europe, to convert the names Pyxicephaloidea, Pyxicephalidae and Pyxicephalinae into phylogenetic nomenclature, following the PhyloCode rules.


  1. Báez, A. M. & Gómez, R. O. (2018) Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil. Journal of systematic Palaeontology, 16: 279–308. <>.

  2. Bittencourt-Silva, G. B., Conradie, W., Siu-Ting, K., Tolley, K., Channing, A., Cunningham, M., Farooq, H., Menegon, M. & Loader, S. (2016) The phylogenetic position and diversity of the enigmatic mongrel frog Nothophryne Poynton, 1963 (Amphibia, Anura). Molecular Phylogenetics and Evolution, 99: 89–102. <>.

  3. Bonaparte, C. (1850) Conspectus Systematum. Herpetologiae et Amphibiologiae. Leiden (Brill): 1 pl.

  4. Bossuyt, F., Brown, R. M., Hillis, D. M., Cannatella, D. C. & Milinkovitch, M. C. (2006) Phylogeny and biogeography of a cosmopolitan frog radiation: Late Cretaceous diversification resulted in continent-scale endemism in the family Ranidae. Systematic Biology, 55: 579–594. <>.

  5. Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. Second edition. London (Taylor & Francis): 1‒530.

  6. Boulenger, G. A. (1887) Descriptions of new reptiles and batrachians in the British Museum (Natural history), part III. Annals and Magazine of natural History, (5), 20 (115): 50‒53. <>.

  7. Boulenger, G. A. (1905). Descriptions of new West-African frogs of the genera Petropedetes and Bulua. Annals and Magazine of natural History, (7), 15 (87): 281‒283. <>.

  8. Boulenger, G. A. (1906) Descriptions of new batrachians discovered by Mr. G. L. Bates in South Cameroon. Annals and Magazine of natural History, (7), 17 (7): 317‒323. <>.

  9. Boulenger, G. A. (1917) Sur la conformation des phalangettes chez certaines grenouilles d’Afrique. Compte rendu hebdomadaire des Séances de l’Académie des Sciences, Paris, 165: 989–990. <>.

  10. Boulenger, G. A. (1918) Aperçu des principes qui doivent régir la classification des espèces du genre Rana. Bulletin de la Société zoologique de France. 43: 111–121. <>.

  11. Boulenger, G. A. (1920a) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum, 20: 1–226. <>.

  12. Boulenger, G. A. (1920b) A monograph of the American frogs of the genus Rana. Proceeding of the American Academy of Arts and Sciences, 55 (9): 413–480. <>.

  13. Branch, W. R. (1976) Two exceptional food records for the African Bullfrog, Pyxicephalus adspersus (Amphibia, Anura, Ranidae). Journal of Herpetology, 10 (3): 266–268. <>.

  14. Cai, Y.-Y., Shen, S.-Q., Lu, L.-X., Storey, K. B., Yu, D.-N. & Zhang, J.-Y. (2019) The complete mitochondrial genome of Pyxicephalus adspersus: high gene rearrangement and phylogenetics of one of the world’s largest frogs. PeerJ, 7 [e7532]. <>.

  15. Cantino, P. D. & Queiroz, K. de (2020) International Code of Phylogenetic Nomenclature (PhyloCode): a phylogenetic code of biological nomenclature. Boca Raton, Florida (CRC Press): 1–189. <>.

  16. Channing, A. (2001) Amphibians of central and southern Africa. Ithaca, NY (Cornell University Press): 1–496. <>.

  17. Channing, A. & Boycott, R. C. (1989) A new frog genus and species from the mountains of the southwestern Cape, South Africa (Anura: Ranidae). Copeia, 1989 (2): 467–471. <>.

  18. Channing, A. & Rödel M.-O. (2019) Field guide to the frogs & other amphibians of Africa. Cape Town, South Africa (Struik Nature): 1–395.

  19. Clarke, B. T. (1981) Comparative osteology and evolutionary relationships in the African Raninae (Anura Ranidae). Monitore zoologico italiano, (n.s.), suppl. 15: 285–331. <>.

  20. De Stefano, G. (1903) I sauri del Quercy appartenenti alla colleztione Rossignol. Atti della Societa italiana di Scienze naturali, 4: 382–418.

  21. Dubois, A. (1981) Liste des genres et sous-genres nominaux de Ranoidea (Amphibiens Anoures) du monde, avec identification de leurs espèces-types: conséquences nomenclaturales. Monitore zoologico italiano, (n.s.), suppl. 15: 225–284. <>.

  22. Dubois, A. (1982) Notes sur la classification des Ranidae (Amphibiens, Anoures). Bulletin mensuel de la Société linnéenne de Lyon, 61 (10): 305–352. <>.

  23. Dubois, A. (1983) Classification et nomenclature supragénérique des Amphibiens Anoures. Bulletin mensuel de la Société linnéenne de Lyon, 52: 270–276. <>.

  24. Dubois, A. (1987) Miscellanaea taxinomica batrachologica (I). Alytes, 5: 7–95.

  25. Dubois, A. (1992) Notes sur la classification des Ranidae (Amphibiens, Anoures). Bulletin mensuel de la Société linnéenne de Lyon, 61: 305–352. <>.

  26. Dubois, A. (1994) Phrynobatrachinae Laurent, 1941 (Amphibia, Anura): proposed conservation. Bulletin of zoological Nomenclature, 51 (3): 240–246. <>.

  27. Dubois, A. (2005) Amphibia Mundi. 1.1. An ergotaxonomy of recent amphibians. Alytes, 23: 1–24.

  28. Dubois, A., Ohler, A. & Pyron, R. A. (2021) New concepts and methods for phylogenetic taxonomy and nomenclature in zoology, exemplified by a new ranked cladonomy of recent Amphibians (Lissamphibia). Megataxa, 5 (1): 1–738. <>.

  29. Duméril, A. H. A. (1856) Note sur les Reptiles du Gabon. Revue et Magasin de Zoologie pure et appliquée, 2: 553–562.

  30. Duméril, A. H. A. & Bibron, G. (1841) Erpétologie générale ou Histoire naturelle complète des Reptiles. Volume 8. Paris (Roret): 1–792.

  31. Feng, Y.-J., Blackburn, D. C., Liang, D., Hillis, D. M., Wake, D. B., Cannatella D. C. & Zhang, P. (2017) Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the national Academy of Sciences, 114: E5864–E5870. <>.

  32. Filhol, H. 1877. Recherches sur les phosphorites du Quercy: étude des fossiles qu’on y rencontre et spécialement des Mammifères. Paris (Librairie de l’Académie de médecine): 1‒690.

  33. Frazão, A., Silva, H. R. da & Russo, C. A. de M. (2015) The Gondwana breakup and the history of the Atlantic and Indian Oceans unveils two new clades for early neobatrachian diversification. PLoS One, 10 [e0143926]. <>.

  34. Frost, D. R., Grant, T., Faivovich, J., Bain, R. H.,  Haas, A., Haddad, C. F. B., de Sá, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. (2006) The Amphibian Tree of Life. Bulletin of the American Museum of natural History, 297: 1–291. <[0001:TATOL]2.0.CO;2>.

  35. Gardner, J. D. & Rage, J.-C. (2016) The fossil record of lissamphibians from Africa, Madagascar, and the Arabian Plate. Palaeobiodiversity and Palaeoenvironments, 96: 169–220. <>.

  36. Hewitt, J. (1919) Anhydrophryne rattrayi, a remarkable new frog from Cape Colony. Records of the Albany Museum, Grahamstown, 3: 182–189.

  37. Hewitt, J. (1925) On some new species of reptiles and amphibians from South Africa. Records of the Albany Museum, Grahamstown, 3: 343–368.

  38. Hewitt, J. (1926) Descriptions of new and little-known lizards and batrachians from South Africa. Annals of the South African Museum, 20: 413–431.

  39. Hewitt, J. & Methuen, P. A. (1912) Descriptions of some new Batrachia and Lacertilia from South Africa. Transactions of the royal Society of South Africa, 3: 107–111. <>.

  40. Jetz, W. & Pyron, R. A. (2018) The interplay of past diversification and evolutionary isolation with present imperilment across the amphibian tree of life. Nature, Ecology & Evolution, 2: 850–858. <>.

  41. Laloy, F., Rage, J. C., Evans, S. E., Boistel, R., Lenoir, N. & Laurin, M. (2013) A re-interpretation of the Eocene anuran Thaumastosaurus based on microCT examination of a ‘mummified’ specimen. PLoS One, 8 [e74874]. <>.

  42. Largen, M. J. (1991) A new genus and species of petropedetine frog (Amphibia Anura Ranidae) from high altitude in the mountains of Ethiopia. Tropical Zoology, 4: 139–152. <>.

  43. Laurent, R. F. (1941) Contribution à l’ostéologie et à la systématique des Ranides africains. Première note. Revue de Zoologie et de Botanique africaines, 34, ‘1940’: 74–96.

  44. Laurent, R. F. (1946) Mises au point dans la taxonomie des Ranides. Revue de Zoologie et de Botanique africaines, 39: 336–338.

  45. Laurent, R. F. (1953) Reptiles et Batraciens récemment parvenus au Musée royal du Congo belge. Bulletin du Cercle zoologique congolais, 21: 21–29.

  46. Laurin, M., Arntzen, J. W., Báez, A. M., Bauer, A. M., Damiani, R., Evans, S. E. et al. (2020a) Amphibia. In: K. de Queiroz, P. D. Cantino & J. A. Gauthier (ed.), Phylonyms: an implementation of PhyloCode, Boca Raton, Florida (CRC Press): 765–771.

  47. Laurin, M., Arntzen, J. W., Báez, A. M., Bauer, A. M., Damiani, R., Evans, S. E. et al. (2020b) Lissamphibia. In: K. de Queiroz, P. D. Cantino & J. A. Gauthier (ed.), Phylonyms: an implementation of PhyloCode, Boca Raton, Florida (CRC Press): 773–778.

  48. Lemierre, A., Folie, A., Bailon, S., Robin N. & Laurin, M. (2021) From toad to frog, a CT-based reconsideration of Bufo servatus, an Eocene anuran mummy from Quercy (France). Journal of Vertebrate Paleontology, 41 (3) [e1989694].

  49. Loveridge, J. P. (1936) Scientific results of an expedition to rain forest regions in eastern Africa VII. Amphibians. Bulletin of the Museum of comparative Zoology, 79: 369–430.

  50. Noble, G. K. (1931) The biology of the Amphibia. New York & London (McGraw-Hill Book Company): 1–600. <>.

  51. Okeyo, D. O., Kandjenogo, L. & Kashea, M. M. (2015) Harvesting and consumption of the giant African bullfrog, a delicacy in northern Namibia. In: K. C. Chinsembu, A. Cheikhyoussef, D. Mumbengegwi, M. Kandawa-Schilz, C. D. Kasanda, & L. Kazembe (ed.), Indigenous knowledge of Namibia, Windhoek, Namibia (UNAM Press): 205–219. <>.

  52. Parry, R. (1982) A revision of southern African Pyxicephalus Tschudi (Anura: Ranidae). Annals of the Natal Museum, 25: 281–292.

  53. Pallas, P. S. (1771) Reise durch verschiedene Provinzen des Russichen Reichs. Theil 1. Saint-Petersburg (Gedruckt bey der Kayserlichen Academie der Wissenschaften): 1–504.

  54. Poynton, J. C. (1963) Descriptions of southern African amphibians. Annals of the Natal Museum, 15: 319–332.

  55. Poynton, J. C. & Broadley, G. (1985) Amphibia Zambesiaca 2. Ranidae. Annals of the Natal Museum, 27: 115–181.

  56. Procter, M. J. B. (1919) On the skull and affinities of Rana subsigillata A. Dum. Proceedings of the zoological Society of London, 89: 21–27. <>.

  57. Pyron, R. A. & Wiens, J. J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61: 543–583. <>.

  58. Queiroz, K. de, Cantino, P. D. & Gauthier, J. A. (2020) Phylonyms: a companion to the PhyloCode. Boca Raton, Florida (CRC Press): 1–1352. <>.

  59. Rage, J.-C., Adaci, M., Bensalah, M., Mahboubi, M., Marivaux, L., Mebrouk, F. & Tabuce, R. (2021) Latest early-middle Eocene deposits of Algeria (Glib Zegdou, HGL50) yield the richest and most diverse fauna of amphibians and squamate reptiles from the Palaeogene of Africa. Paleovertebrata, 44 (1) [e1]. <>.

  60. Robbins, L. H., Murphy, M. L., Stevens, N. J., Brook, G. A., Ivester, A. H., Haberyan, K. A., Klein, R. G., Milo, R., Stewart, K. M., Matthiesen, D. G., & Winkler, A. J. (1996) Paleoenvironment and archaeology of Drotsky’s Cave: western Kalahari Desert, Botswana. Journal of archaeological Science, 23: 7–22. <>.

  61. Robbins, L. H., Campbell, A. C., Murphy, M. L., Brook, G. A., Mabuse, A. A., Hitchcock, R. K., Babutsi, G., Mmolowa, M., Stewart, K.M., Steele, T. E., Klein, R. G. & Appleton, C. C. (2009). Mogapelwa: archaeology, palaeoenvironment and oral traditions at Lake Ngami, Botswana. South African archaeological Bulletin, 64: 13–32.

  62. Roček, Z. & Lamaud P. (1995) Thaumastosaurus bottii De Stefano, 1903, an anuran with Gondwanan affinities from the Eocene of Europe. Journal of Vertebrate Paleontology, 15: 506–515. <>.

  63. Rödel, M.-O. (2000) Herpetofauna of West Africa. Volume 1. Amphibians of the West African savanna. Frankfurt/Main (Edition Chimaira): 1–335.

  64. Scott, E. (2005) A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data. Cladistics, 21: 507–574. <>.

  65. Smith, A. (1849) Illustrations of the zoology of South Africa, consisting chiefly of figures and descriptions of the objects of natural history collected during an expedition into the interior of South Africa in the years 1834, 1835 and 1836... Reptilia. London (Smith, Elder & Co.): appendix 1–28. <>.

  66. Trueb, L. (1973) Bones, frogs and evolution. In: J. L. Vial (ed.), Evolutionary biology of the Anurans, Contemporary research on major problems, Columbia (University of Missouri Press): 65–133.

  67. Tschudi, J. J. (1838) Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Mémoires de la Société des Sciences naturelles de Neuchâtel, 2: 1–100. <>.

  68. Van der Meijden, A., Crottini, A., Tarrant, J., Turner, A. & Vences, M. (2011) Multi-locus phylogeny and evolution of reproductive modes in the Pyxicephalidae, an African endemic clade of frogs. African Journal of Herpetology, 60: 1–12. <>.

  69. Van der Meijden, A., Vences, M., Hoegg, S. & Meyer, A. (2005) A previously unrecognized radiation of ranid frogs in Southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 37: 674–685. <>.

  70. Van Dijk, D. E. (1966) Systematic and field keys to the families, genera and described species of southern African anuran tadpoles. Annals of the Natal Museum, 18: 231–286.

  71. Vasilyan, D. (2018) Eocene Western European endemic genus Thaumastosaurus: new insights into the question “Are the Ranidae known prior to the Oligocene?” PeerJ, 6 [e5511]. <>.

  72. Wake, D. B. (2020) Caudata. In: K. de Queiroz, P. D. Cantino & J. A. Gauthier (ed.), Phylonyms: an implementation of PhyloCode. Boca Raton, Florida (CRC Press): 785–787.

  73. Wake, M. H. (2020) Gymnophiona. In: K. de Queiroz, P. D. Cantino & J. A. Gauthier (ed.), Phylonyms: an implementation of PhyloCode. Boca Raton, Florida (CRC Press): 779–783.

  74. Yuan, Z.-Y., Zhang, B.-L., Raxworthy, C. J., Weisrock, D. W., Hime, P. M., Jin, J.-Q., Lemmon, E. M., Lemmon, A. R., Holland, S. D., Kortyna, M. L., Zhou, W.-W., Peng, M.-S., Che, J., & Prendini, E. (2019) Natatanuran frogs used the Indian Plate to step-stone disperse and radiate across the Indian Ocean. National Science Review, 6: 10–14. <>.