Skip to main content Skip to main navigation menu Skip to site footer
Published: 2021-12-17

Barton and the forgotten description of Salamandra horrida (Amphibia, Urodela): story of a rediscovery

Association RACINE, 5 allée des cygnes, 35750 Saint Maugan, France
Penclen, 56420 Plumelec, France
Cryptobranchidae Cryptobranchus Cryptobranchus horridus hellbender logonymic lists new synonymy first revisor action airesy subsequent monophory lineages


Cryptobranchus alleganiensis (Sonnini & Latreille, 1801b) was long considered the only species of the genus Cryptobranchus Leuckart, 1821. In this genus, Sabatino & Routman (2009) found eight genetic independent unities isolated from each other, which they treated as Management Units. Later, Hime (2017) recovered five strongly supported species-level lineages within this genus, corresponding more or less to the lineages of Sabatino & Routman (2009). We herein resurrect a valid nomen for one of the lineages of Hime (2017) (Ohio/Allegheny/Susquehanna lineage), the most broadly distributed of the five, which should be named Cryptobranchus horridus (Barton, 1807a). We relate the story of the rediscovery of two forgotten publications of Barton (1807a‒b) preceding that of 1808 so far recognized as the original description of this taxon. We discuss the problem of the onymotope of Cryptobranchus alleganiensis and present logonomic lists for the genus Cryptobranchus and all the taxa described. The nucleospecies of the genus Cryptobranchus is Salamandra horrida Barton, 1807a which is no longer a synonym of Salamandra alleganiensis Sonnini & Latreille, 1801b. The updating of the logonymic lists revealed the forgotten designation of Barnes (1828) for the genus Protonopsis Le Conte, 1824, a synonym of Cryptobranchus. The following nomina can be allocated to three of the five lineages identified by Hime (2017): Cryptobranchus alleganiensis, Cryptobranchus horridus and Cryptobranchus bishopi. Two other lineages require further studies.


  1. Anonymous [International Commission on Zoological Nomenclature] (1926) Opinion 92. Sixteen generic names of Pisces, Amphibia, and Reptilia placed in the Official List of Generic Names. In: Opinions rendered by the International Commission on Zoological Nomenclature, Opinions 91 to 97, in: Smithsonian miscellaneous Collections, 73 (4): 3–4.

  2. Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. ‘Fourth edition’. London (International Trust for zoological Nomenclature): i–xxix + 1–306.

  3. Adler, K. (2012) Contributions to the history of herpetology. Volume 3. Contributions to Herpetology, New York (Society for the Study of Amphibians and Reptiles), 29: 1–564.

  4. Barnes, D. H. (1826) An arrangement of the genera of batracian animals, with a description of the more remarkable species; including a monograph of the doubtful reptils. American Journal of Science and Arts, 11: 268–297.

  5. Barnes, D. H. (1828) Note on the doubtful reptils. American Journal of Science and Arts, 13: 66–70.

  6. Barton, [B. S.] (1807a) Amphibiology. Philadelphia medical and physical Journal, Supplement 2 (2): 195–197.


  8. Barton, [B. S.] (1807b) A discourse on some of the principal desiderata in natural history, and on the best means of promoting the study of this science, in the United-States. Philadelphia (Denham & Town): 1–90.

  9. Barton, [B. S.] (1808) Some Account of the Siren lacertina, and other species of the same genus of amphibious animals. In a Letter from Professor Barton, of Philadelphia, to Mr. John Gottlob Schneider, of Saxony: 1–36, 1 pl.

  10. Barton, [B. S.] (1814) A memoir concerning an animal of the class of Reptilia, or Amphibia, which is known, in the United-States, by the names of alligator and hell-bender. Philadelphia (Griggs and Dickinson), ‘1812’: i–iv + 5–26, 1 pl.

  11. Barton, [B. S.] (1821) Some account of the Siren lacertina, and other species of the same genus of amphibious animals in a letter from professor Barton, of Philadelphia to Mr. John Gottlob Schneider, of Saxony. Philadephia (Jesper Harding): 1–32, 1 pl. <>.

  12. Bauer, A. M., Good, D. A. & Günther, R. (1993) An annotated type catalogue of the caecilians and salamanders (Amphibia: Gymnophiona and Caudata) in the Zoological Museum, Berlin. Mitteilungen aus dem zoologischen Museum in Berlin, 69 (2): 285–306. <>.

  13. Boïé, [H.] (1824) Extrait d’une lettre du Dr. J. C. van Hasselt au Prof. van Swinderen, sur les Mollusques de Java. Bulletin des Sciences naturelles et de Géologie, 3: 237–245.

  14. Bonaparte, C. L. (1839) Euproctus platycephalus, Euprotto del Rusconi. In: Iconografia della fauna italica per le quattro classi degli animali vertebrati, Volume 2, Fasciculo 26, Roma (Salviucci): [259–266], pl. [85].

  15. Bosc, [L. A. G.] (1803) Salamandre. Nouveau dictionnaire d’histoire naturelle. Paris (Déterville), 20: 41–50, pl. 12.

  16. Boulenger, G. A. (1882) Catalogue of the Batrachia Gradientia s. Caudata and Batrachia Apoda in the collection of the British Museum. Second edition. London (Trustees of the British Museum, Natural History): i–ix + 1–127, pl. 1–9.

  17. Brown, A. E. (1908) Generic types of Nearctic Reptilia and Amphibia. Proceedings of the Academy of natural Sciences of Philadelphia, 60: 112–127.

  18. Collins, J. T. (1991) Viewpoint: a new taxonomic arrangement for some North American Amphibians and Reptiles. Herpetological Review, 22 (2): 42–43.

  19. Cope, E. D. (1889) The Batrachia of North America. Bulletin of the United States national Museum, 34: 1–525, pl. 1–86.

  20. Cuvier, G. (1816) Le règne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Tome 2, contenant les Reptiles, les Poissons, les Mollusques et les Annélides. Paris (Deterville), ‘1817’: i–xviii + 1–532.

  21. Daudin, F. M. (1803) Histoire naturelle, générale et particulière, des Reptiles. Volume 8. Paris (F. Dufart): 1–439, pl. 93–100.

  22. Davis, N. S. & Rice, F. L. (1883) List of Batrachia and Reptilia of Illinois. Bulletin of the Chicago Academy of Sciences, 1 (3): 25–32. <>.

  23. Dubois, A. (2000) Synonymies and related lists in zoology: general proposals, with examples in herpetology. Dumerilia, 4 (2): 33–98.

  24. Dubois, A. (2005a) Propositions pour l’incorporation des nomina de taxons de rang supérieur dans le Code international de nomenclature zoologique. In: A. Dubois, O. Poncy, V. Malécot & N. Léger (ed.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23: 73–96.

  25. Dubois, A. (2005b) Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code of Zoological Nomenclature. 1. Some general questions, concepts and terms of biological nomenclature. Zoosystema, 27 (2): 365–426.

  26. Dubois, A. (2006) Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code of Zoological Nomenclature. 2. The proposed Rules and their rationale. Zoosystema, 28 (1): 165–258.

  27. Dubois, A. (2010) Retroactive changes should be introduced in the Code only with great care: problems related to the spellings of nomina. Zootaxa, 2426: 1–42. <>.

  28. Dubois, A. (2011) The International Code of Zoological Nomenclature must be drastically improved before it is too late. Bionomina, 2: 1–104. <>.

  29. Dubois, A. (2013) Zygoidy, a new nomenclatural concept. Bionomina, 6: 1–25. <>.

  30. Dubois, A. (2015) Zoological nomina in the century of extinctions: new proposals. Bionomina, 8: 11–53. <>.

  31. Dubois, A. (2017) A few problems in the generic nomenclature of insects and amphibians, with recommendations for the publication of new generic nomina in zootaxonomy and comments on taxonomic and nomenclatural databases and websites. Zootaxa, 4237 (1): 1–16. <>.

  32. Dubois, A. & Aescht, E. (ed.) (2019) LZC Session 28. The Principles of the Zoocode. 10. The Principle of Synonymy. Dumerilia, 8: 106–109.

  33. 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. <>.


  35. Duméril, A.-M.-C., Bibron, G. & Duméril, A. (1854) Erpétologie générale ou histoire naturelle complète des Reptiles. Tome 9. Paris (Roret): i–xx + 1–440. <>.

  36. Dundee, H. A. (1972) Cryptobranchus, and C. alleganiensis. Catalogue of American Amphibians and Reptiles, 101: 1–4.

  37. Eichwald, E. (1831) Zoologia specialis quam expositis animalis tum vivis, tum fossilibus potossimum Rossiae in universum, et Poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi habendarum. Pars posterior specialem expositionem spondylozoorum continensis. Vilniae (Josephi Zawadski): 1–404, pl. 1.

  38. Fitzinger, L. I. (1826) Neue Classification der Reptilien. Wien (Heubner): i–viii + 1–66, 1 tab.

  39. Fitzinger, L. I. (1843) Systema Reptilium. Fasc. 1. Amblyglossae. Vindobonae (Braumüller & Seidel): 1–106 + i–ix.

  40. Fouquette, M. J., Jr. & Dubois, A. (2014) A checklist of North American Amphibians and Reptiles. Volume 1. Amphibians. USA (XLibris): 1–613.

  41. Frétey, T., Dewynter, M. & Ohler, A. (2018) Onymotopes in zoological nomenclature: some additional terms, with fixation of a lectonymotope for Xenopus petersii Bocage, 1895 (Amphibia, Anura). Bionomina, 13: 37–50. <>.

  42. Frétey, T. & Dubois, A. (2019) The authorship and date of five generic nomina of Squamata and Amphibia published by Cuvier (1816, 1829), with recourse to and comments on twenty Articles of the Code. Bionomina, 15: 1–36. <>.

  43. Gray, J. E. (1825) A synopsis of the genera of Reptiles and Amphibia, with a description of some new species. Annals of Philosophy, (2), 10: 193–217.

  44. Gray, J. E. (1831) A synopsis of the species of the Class Reptilia. In: E. Griffith, The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, &c. &c. &c. with additional descriptions of all the species hitherto named, and of many not before noticed. London (Whittaker, Treacher & Co.), 9 (2): 1–110.

  45. Gray, J. E. (1850) Catalogue of the specimens of Amphibia in the collection of the British Museum. Part 2. Batrachia Gradientia, etc. London (Spottiswoodes & Shaw): 1–72, pl. 3–4.

  46. Griffith, E. (1835) A classified index and synopsis of the Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, &c. &c. &c. with supplementary additions to each order. London (Whittaker & Co.): i–cxix + 1–328.

  47. Griffith, E. & Pidgeon, E. (1831) The Class Reptilia arranged by the Baron Cuvier with specific descriptions. In: E. Griffith, The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, &c. &c. &c. with additional descriptions of all the species hitherto named, and of many not before noticed. London (Whittaker, Treacher & Co.), 9 (1): [i–ii] + 1–481, 55 pl.

  48. Grobman, A. B. (1943) Notes on salamanders with the description of a new species of Cryptobranchus. Occasional Papers of the Museum of Zoology, University of Michigan, 470: 1–12.

  49. Guérin-Méneville, F. E. (1838) Reptiles. In: Iconographie du Règne Animal de G. Cuvier ou Représentation d’après nature de l’une des espèces les plus remarquables et souvent non encore figurées, de chaque genre d’animaux, avec un texte descriptif mis au courant de la science. Volume 3. Paris (J. B. Ballière): 1–23, errata.

  50. Harlan, R. (1825a) Observations on the genus Salamandra, with the anatomy of the Salamandra gigantea (Barton) or S. Alleghaniensis (Michaux) and two new genera proposed. Annals of the Lyceum of natural History of New-York, 1 (2): 222–234, pl. 16–18. <>.

  51. Harlan, R. (1825b) Note to a paper entitled observations on the genus Salamandra. Annals of the Lyceum of natural History of New-York, 1 (2): 270–271. <>.

  52. Harlan, R. (1827). Genera of North American Reptilia, and a synopsis of the species. Journal of the Academy of natural Sciences of Philadelphia, 5 (2): 317–372.

  53. Harlan, R. (1835) Medical and physical researches: or original memoirs in medicine, surgery, physiology, geology, zoology, and comparative anatomy. Philadelphia (Lydia R. Bailey): i–xxxix + 9–653, 40 pl. <>.

  54. Harper, F. A. (1940) Some works of Bartram, Daudin, Latreille, and Sonnini, and their bearing upon North American herpetological nomenclature. American Midland Naturalist, 23: 692–723. <>.

  55. Hemming, F. (1956) Direction 57. Addition to the Official List of Specific Names in Zoology (a) of the specific names of forty-seven species belonging to the classes Cyclostomata, Pisces, Amphibia and Reptilia, each of which is the type species of a genus, the name of which was placed on the Official List of Generic Names in Zoology in the period up to the end of 1936 and (b) of the specific name of one species of the class Amphibia which is currently treated as senior subjective synonym of the name of such a species. Opinions and Declarations rendered by the International Commission on Zoological Nomenclature, 1D (18): 365–388.

  56. Hime, P. M. (2017) Genomic perspectives on Amphibian evolution across multiple phylogenetic scales. Thesis, Lexington (University of Kentucky): i–xv + 1–257.

  57. Holbrook, J. E. (1842) North American herpetology; or, a description of the Reptiles inhabiting the United States. Volume 5. Philadelphia (J. Dobson): i–iv + 5–118. <>.


  59. Holman, J. A. (1977) The Pleistocene (Kansan) herpetofauna of Cumberland Cave, Maryland. Annals of the Carnegie Museum, 46 (11): 157–172.

  60. James, E. (1823) Account of an expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and ’20. Philadelphia (H. C. Garey & I. Lea): [i–iii] + 1–7 + 1–503. <>.

  61. Knauer, F. K. (1878) Naturgeschichte der Lurche. (Amphibiologie). Wien (Pichler’s Witwe & Sohn): i–xx + 1–340, 5 pl.

  62. Le Conte, J. E. (1824) Description of a new species of Siren, with some observations on animals of a similar nature. Annals of the Lyceum of natural History of New-York, 1 (1): 52–58. <>.

  63. Leuckart, S. (1821) Einiges über die Fischartigen Amphibien. Isis von Oken, Litterarischer Anzeiger, 1821: 257–265, pl. 5.

  64. Leuckart, F. S. (1840) Einige zoologische Bemerkungen. 2. Ueber das Gen. Cryptobranchus. Neue Notizen aus dem Gebiete der Natur- und Heilkunde, 13 (2): 19–20.

  65. Merrem, B. (1820) Versuch eines Systems der Amphibien. Tentamen systematis Amphibiorum. Marburg (Iohann Christian Krieger): [i–xv + 1–191] × 2, 1 pl. <>.

  66. Michaux, A. (1804) Voyage à l’ouest des monts Alléghanys, dans les états de l’Ohio, du Kentucky et du Tennessee et retour à Charleston par les Hautes-Carolines. Paris (Crapelet): i–vi + 1–312, 1 map. <>.

  67. Michaux, F. A. (1805) Travels to the westward of the Allegany Mountains, in the States of the Ohio, Kentucky, and Tennessee and return to Charlestown, through the Upper Carolinas. London (J. Mawman): i–xvi +1–350, 1 map.

  68. Michaux, A. (1889) Portions of the journal of Andre Michaux, botanist, written during his travels in the United States and Canada, 1785 to 1796. With an introduction and explanatory notes, by C. S. Sargent. Proceedings of the American philosophical Society, 26 (129): 1–145.

  69. Morse, M. (1901) Ohio Batrachia in the Zoological Museum of the O. S. U. The Ohio Naturalist, 1 (7): 114–115.

  70. Nickerson, M. A. & Mays, C. E. (1973) The hellbenders: North American ‘giant salamanders’. Publications in Biology and Geology, Milwaukee public Museum, 1: i–vii + 1–106.

  71. Oppel, M. (1811) Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben. München (Lindauer): i–xii + 1–87. <>.

  72. Raffaëlli, J. (2013) Les Urodèles du Monde. Deuxième édition. Plumelec (Penclen edition): 1–472.

  73. Rafinesque, C. S. (1819) Prodrome de 70 nouveaux genres d’animaux découverts dans l’intérieur des États-Unis d’Amérique, durant l’année 1818. Journal de Physique, de Chimie, d’Histoire naturelle et des Arts, 88: 417–429.

  74. Rafinesque, C. S. (1822) On two new salamanders of Kentucky. Kentucky Gazette, (n.s.), 1 (9): 3.

  75. Rafinesque, C. S. (1832) On 3 new water salamanders of Kentucky. Atlantic Journal and Friend of Knowledge, 1: 121.

  76. Sabatino, S. J. & Routman, E. J. (2009) Phylogeography and conservation genetics of the hellbender salamander (Cryptobranchus alleganiensis). Conservation Genetics, 10: 1235–1246. <>.

  77. Schlegel, H. (1858) Handleiding tot de Beoefening der Dierkunde. Breda (Koninklijke Militaire Akademie), 2: i–xx + 1 + i–xx + 1–630. <>.

  78. Schmidt, K. P. (1953) A check list of North American Amphibians and Reptiles. Sixth edition. American Society of Ichthyologists and Herpetologists and University of Chicago Press: i–viii + 1–280.

  79. Simpson, G. G. (1940) Types in modern taxonomy. American Journal of Science, 238: 413–431. <>.

  80. Smith, H. M. & Pérez-Higareda, G. (1986) Nomenclatural name-forms. Systematic Zoology, 35 (3): 421–422. <>.

  81. Smith, J. E. (1821) A selection of the correspondence of Linnaeus and other naturalists from the original manuscripts. Volume 1. London(Longman, Hurst, Rees, Orme & Brown): i–xiv + 1–605.

  82. Smith, W. H. (1877) The Tailed Amphibians, including the Caecilians. A thesis presented to the Faculty of Michigan University. Detroit (Herald Publishing House): 1–158 + 6 tab. <>.

  83. Sonnini, C. S. & Latreille, P. A. (1801a) [An X] Histoire naturelle des Reptiles, avec figures dessinées d’après nature. Tome 2. Première partie. Quadrupèdes et bipèdes ovipares. Paris (Deterville): 1–332, 20 pl. <>.

  84. Sonnini, C. S. & Latreille, P. A. (1801b) [An X] Histoire naturelle des Reptiles, avec figures dessinées d’après nature. Tome 4. Seconde partie. Serpens. Paris (Deterville): 1–410, 14 pl. <>.

  85. Stejneger, L. & Barbour, T. (1939) A check list of North American Amphibians and Reptiles. Fourth edition. Cambridge (Harvard University Press): i–xvi + 1–207. <>.

  86. Stuart, S. N., Hoffmann, M., Chanson, J. S., Cox, N. A., Berridge, R. J., Ramani, P. & Young, B. E. (ed.) (2008) Threatened Amphibians of the World. Barcelona (Lynx Edicions): i–xv + 1–758.

  87. Tschudi, J. J., von (1838) Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Neuchâtel (Petitpierre): 1–98, pl. 1–6 + [i–ii]. <>.

  88. Van der Hoeven, J. (1833) Handboek der Dierkunde, of Grondbeginsels der Natuurlijke Geschiedenis van het Dierenrijk. Volume 2. Amsterdam (C. G. Sulpke): i–x + i–v + 1–698 + 1–10.


  90. Van der Hoeven, J. (1838) Iets over den grooten zoogenoemden den Salamander von Japan. Tidjschrift voor natuurlijke Geschiedenis en Physiologie, ‘1837’, 4: 375–386, 2 pl.

  91. Van Hasselt, J. C. (1824) Uittreksel uit eenen Brief van Dr. J. C. van Hasselt, aan Prof. van Swinderen. Algemeene Konst-en Letter-Bode, 2: 20–24 + 34–39 + 54–55.

  92. Wagler, J. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vögel. München, Stuttgart & Tübingen (Cotta): i–vi + 1–354. <>.

  93. Waite, F. C. (1907) Specific name of Necturus maculosus. American Naturalist, 41: 23–30. <>.

  94. Wellborn, V. (1936) Beschreibung eines neuen Molches der Gattung Cryptobranchus. Zoologischer Anzeiger, 114: 63–64.

  95. Yarrow, H. C. (1882) Check list of North American Reptilia and Batrachia with catalogue of specimens in U. S. National Museum. Bulletin of the United States national Museum, 24: i–viii + 1–249. <>.