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Type: Article
Published: 2022-05-10
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A nomenclatural and taxonomic review of the salamanders (Urodela) from Holbrook’s North American Herpetology

Department of Biological Sciences, The George Washington University, 2023 G St. NW, Washington, DC 20052
Department of Natural Sciences, Nash Community College, Rocky Mount, NC 27804
Amphibia Holbrook North American Herpetology salamanders Urodela type specimen

Abstract

John Edwards Holbrook published North American Herpetology in 11 volumes from 1836–1842, authoring the first accounts of numerous amphibians and reptiles from the eastern and central United States, including 32 salamanders (Urodela). We reviewed these and located 51 extant salamander specimens from Holbrook in the Academy of Natural Sciences (Philadelphia), Museum of Comparative Zoology (Cambridge), and Muséum national d’Histoire naturelle (Paris), six of which are types. We identified four other specimens figured by Holbrook in the MNHN and National Museum of Natural History (Washington), all of which are types from descriptions by other authors. We designate lectotypes for S. porpyhritica Green, 1827 (USNM 3840; reversing neotype MCZ A-35778), Salamandra gutto-lineata Holbrook, 1838a (ANSP 716), S. auriculata Holbrook, 1838b (MNHN-RA 0.4675), S. maculo-quadrata Holbrook, 1840 (ANSP 821), S. granulata De Kay in Holbrook, 1842e (USNM 3981), S. quadridigitata Holbrook, 1842e (ANSP 490; reversing neotype UF 178833), and Plethodon variolosum Duméril, Bibron, and Duméril, 1854 (MNHN-RA 0.4666). Allocation of S. auriculata Holbrook, 1838b, S. “Haldemani” Holbrook, 1840, and P. variolosum Duméril, Bibron, and Duméril, 1854 is still ambiguous. We consider S. maculo-quadrata Holbrook, 1840 to be a junior subjective synonym of S. fusca Green, 1818; no valid name has ever been applied to Black-bellied Salamanders (Desmognathus sp. “quadramaculatus”) at the species level, and up to five candidate species require new names. Additional discoveries of data and specimens pertaining to Holbrook’s names may remain to be made among his surviving papers and collections.

 

References

  1. Adler, K. (1976) New genera and species described in Holbrook’s “North American Herpetology”. In: Adler, K. (Ed.), Holbrook’s North American Herpetology by John Edwards Holbrook. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, pp. xxix–xlii.
    Adler, K. (1989) Contributions to the history of herpetology: Issued to commemorate the I World Congress of Herpetology, Canterbury 1989. Vol. 1. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 202 pp.
    Anderson, J.D. (1967) Ambystoma maculatum. Catalogue of American Amphibians and Reptiles, 51, 1–4.
    Anderson, W.D. Jr. (2014) The plates in Holbrook’s Ichthyology of South Carolina, 1860. aqua: International Journal of Ichthyology, 20 (4), 173–218.
    Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. Fourth edition. International Trust for zoological Nomenclature, London, xxix + 306 pp. https://doi. org/10.5962/bhl.title.50608.
    Anonymous [International Commission on Zoological Nomenclature] (2007) New applications to the Commission. Bulletin of Zoological Nomenclature, 64, 137–138.
    Anonymous [International Commission on Zoological Nomenclature] (2008) Notice of closure of Cases. Bulletin of Zoological Nomenclature, 65, 329–329. https://doi.org/10.21805/bzn.v65i4.a3
    Bailey, J.R. (1937) Notes on plethodont salamanders of the southeastern United States. Occasional Papers of the Museum of Zoology, University of Michigan, 364, 1–10.
    Baird, S.F. (1850) “1849” Revision of the North American Tailed-Batrachia, with descriptions of new genera and species. Journal of the Academy of Natural Sciences of Philadelphia, 2 (1), 281–294.
    Bakker, F.T., Antonelli, A., Clarke, J.A., Cook, J.A., Edwards, S.V., Ericson, P.G.P., Faurby, S., Ferrand, N., Gelang, M., Gillespie, R.G., Irestedt, M., Lundin, K., Larsson, E., Matos-Maraví, P., Müller, J., Proschwitz, T. von, Roderick, G.K., Schliep, A., Wahlberg, N., Wiedenhoeft, J. & Källersjö, M. (2020) The Global Museum: natural history collections and the future of evolutionary science and public education. PeerJ, 8, e8225. https://doi.org/10.7717/peerj.8225
    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.
    Barnes, D.H. (1827) Note on the doubtful reptils. American Journal of Science and Arts, 13, 66–70.
    Barnhart, J.H. (1917) John Eatton Le Conte. The American Midland Naturalist, 5, 135–138. https://doi.org/10.2307/2992778
    Barton, B.S. (1807) Amphibiology. Philadelphia medical and physical Journal, Supplement, 2, 195–197.
    Barton, B.S. (1804) Some account of a new species of North American lizard. Transactions of the American Philosophical Society, 6 (1), 108–112. https://doi.org/10.2307/1004778
    Bauer, A.M. (2003) On the identity of Lacerta punctata Linnaeus 1758, the type species of the genus Euprepis Wagler 1830, and the generic assignment of Afro-Malagasy skinks. African Journal of Herpetology, 52 (1), 1–7. https://doi.org/10.1080/21564574.2003.9635472
    Beamer, D.A. & Lamb, T. (2008) Dusky salamanders (Desmognathus, Plethodontidae) from the Coastal Plain: Multiple independent lineages and their bearing on the molecular phylogeny of the genus. Molecular Phylogenetics and Evolution, 47, 143–153. https://doi.org/10.1016/j.ympev.2008.01.015
    Beamer, D.A. & Lamb, T. (2020) Towards rectifying limitations on species delineation in dusky salamanders (Desmognathus: Plethodontidae): An ecoregion-drainage sampling grid reveals additional cryptic clades. Zootaxa, 4734 (1), 1–61. https://doi.org/10.11646/zootaxa.4734.1.1
    Bechstein, J.M. (1800) Herrn De la Cepede’s Naturgeschichte der Amphibien oder der eyerlengenden vierfüssigen Thiere und der Schlangen. Eine Fortsetzung von Buffon’s Naturgeschichte aus dem Französischen übersetzt und mit Anmerkungen und Zusätsen versehen. Vol. 2. Industrie Comptoir, Weimar, XX + 552 pp.
    Bell, E.L. (1996) Descriptions of neotypes for Sceloporus undulatus undulatus, the Southern Fence Lizard, and Sceloporus undulatus hyacinthinus, the Northern Fence Lizard, and a lectotype for Sceloporus undulatus garmani, the Northern Prairie Lizard. Bulletin of the Maryland Herpetological Society, 32, 81–103.
    Bell, T. (1839) A History of British Reptiles. John van Voorst, London, xxiv + 142 pp. https://doi.org/10.5962/bhl.title.10835
    Bingham, R.E., Papenfuss, T.J., Lindstrand, L. & Wake, D.B. (2018) Phylogeography and species boundaries in the Hydromantes shastae complex, with description of two new species (Amphibia; Caudata; Plethodontidae). Bulletin of the Museum of Comparative Zoology, 161, 403–427. https://doi.org/10.3099/MCZ42.1
    Bishop, S.C. (1941) Notes on salamanders with descriptions of several new forms. Occasional Papers of the Museum of Zoology, University of Michigan, 451, 1–21.
    Bonnaterre, P.-J. (1789) Tableau Encyclopédique et Méthodique des Trois Règnes de la Nature. Erpétologie. Panckouke, Paris, XXVIII + 71 pp., 35 pls. https://doi.org/10.5962/bhl.title.11660
    Brame, A. (1972) Checklist of living and fossil salamanders of the world. Pasadena, California, 1–387. [unpublished manuscript]
    Brandon, R.A. (1966) Systematics of the salamander genus Gyrinophilus. Illinois Biological Monographs, 35, 1–86. https://doi.org/10.5962/bhl.title.50088
    Brongniart, A. (1800) Essai d’une classification naturelle des reptiles. Seconde Partie. Formation et disposition des genres. Bulletin des Sciences, par la Société Philomathique de Paris, 2, 89–91.
    Bryan, C.S. & Whitehead, A.W. (2014) “An aristocracy of talent”: the South Carolina physician-naturalists and their times. Transactions of the American Clinical and Climatological Association, 125, 313–330.
    Burbrink, F.T., Pyron, R.A., Gehara, M.C., Mckelvy, A.D. & Myers, E.A. (2021) The corrected taxonomic history of the North American ratsnakes (Pantherophis obsoletus complex). Herpetological Review, 53, 537–547.
    Cardarelli, F. (2003) Encyclopaedia of scientific units, weights, and measures: their SI equivalences and origins. Springer, London & New York, xxiv + 848 pp. https://doi.org/10.1007/978-1-4471-0003-4_1
    Carlin, J.L. (1997) Genetic and morphological differentiation between Eurycea longicauda longicauda and E. guttolineata (Caudata: Plethodontidae). Herpetologica, 53, 206–217.
    Catesby, M. (1747) Appendix. In: Catesby, M. (Ed.), The natural history of Carolina, Florida, and the Bahama Islands. “MDCCXXXI-MDCCXLIII.” Vol. 2. Printed at the expence of the author, and sold by W. Innys and R. Manby, at the West End of St. Paul’s, by Mr. Hauksbee, at the Royal Society House, and by the author, at Mr. Bacon’s in Hoxton, London, pp. 1–20.
    Catesby, M. (1754) The natural history of Carolina, Florida, and the Bahama Islands. Volume I. C. Marsh and T. Wilcox, London, vii + 122 pp.
    Catesby M. (1771) The natural history of Carolina, Florida, and the Bahama Islands. Volume II. Benjamin White, London, vii + xliv + 100 pp., 100 pls.
    Cochran, D.M. (1961) Type specimens of reptiles and amphibians in the U.S. National Museum. Bulletin of the United States National Museum, 220, 1–291. https://doi.org/10.5962/bhl.part.26967
    Cope, E.D. (1869) A review of the species of Plethodontidae and Desmognathidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 21, 93–118.
    Cope, E.D. (1889) The Batrachia of North America. Bulletin of the United States National Museum, 34, 1–525. https://doi.org/10.5962/bhl.title.38254
    Cope, E.D. (1859) On the primary divisions of the Salamandridae, with descriptions of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 11, 122–128.
    Cope, E.D. (1867) A review of the species of the Amblystomidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 19, 166–211.
    Cuvier, G.L.C.F.D. (1827) Sur le genre de reptiles batraciens, nomme Amphiuma, et sur une nouvelle espèce de ce genre (Amphiuma tridactylum). Mémoires du Muséum d’Histoire Naturelle, Paris, 14, 1–14.
    Daubenton, L.J.-M. (1782) Encyclopédie Méthodique. Histoire Naturelle. Les Animaux Quadrupèdes Ovipaires, et les Serpens. Tome I. Panckouke, Paris, 691 pp.
    Daudin, F.M. (1803) Histoire naturelle, générale et particulière, des reptiles. Tome Huitième. F. Dufart, Paris, 439 pp.
    De Kay, J.E. (1842) Zoology of New-York, or, The New-York Fauna: Comprising Detailed Descriptions of All the Animals Hitherto Observed within the State of New-York, with Brief Notices of those Occasionally Found Near its Borders, and Accompanied by Appropriate Illustrations. Part III. Reptiles and Amphibia. W. & A. White & J. Visscher, Albany, 98 pp. https://doi.org/10.5962/bhl.title.13735
    Dowling, H.G. (1952) A taxonomic study of the ratsnakes, genus Elaphe Fitzinger, IV: A check list of the American forms. Occasional Papers of the Museum of Zoology, University of Michigan, 541, 1–12.
    Dubois, A., Nemésio, A. & Bour, R. (2014) Primary, secondary and tertiary syntypes and virtual lectotype designation in zoological nomenclature, with comments on the recent designation of a lectotype for Elephas maximus Linnaeus, 1758. Bionomina, 7 (1), 45–64. https://doi.org/10.11646/bionomina.7.1.2
    Duméril, A.M.C., Bibron, G. & Duméril A.H.A. (1854) Erpétologie Genérale, ou, Histoire Naturelle Complète des Reptiles. Tome Neuvième, Première Partie. Librarie Enclyclopedique de Roret, Paris, xx + 440 pp. https://doi.org/10.5962/bhl.title.118797
    Dunn, E.R. (1917) The salamanders of the genera Desmognathus and Leurognathus. Proceedings of the United States National Museum, 53, 393–433. https://doi.org/10.5479/si.00963801.53-2211.393
    Dunn, E.R. (1926) The salamanders of the family Plethodontidae. Smith College, Northampton, Massachusetts, 446 pp.
    Edwards, G. (1771) A catalogue of the animals and plants represented in Catesby’s Natural History of Carolina: with the Linnaean names. In: Catesby, M. (Ed.), The natural history of Carolina, Florida, and the Bahama Islands. Vol. II. Benjamin White, London, pp. 1–2.
    Ferraris, C.J. & Vari, R.P. (1992) Catalog of type specimens of recent fishes in the National Museum of Natural History, Smithsonian Institution. Smithsonian Contributions to Zoology, 535, 1–52. https://doi.org/10.5479/si.00810282.535
    Fitzinger, L.J.F.J. (1826) Neue Classification der Reptilien nach ihren natürlichen Verwandtschaften: nebst einer verwandtschafts-tafel und einem Verzeichnisse der Reptilien-sammlung des K. K. zoologischen Museum’s zu Wien. J.G. Heubner, Wien, 61 pp. https://doi.org/10.5962/bhl.title.4683
    Fleming, J. (1822) The philosophy of zoology: or, a general view of the structure, functions, and classification of animals. Volume Two. A. Constable, Edinburgh, 618 pp. https://doi.org/10.1037/11823-000
    Fouquette, M.J. & Dubois, A. (2014) A checklist of North American amphibians and reptiles: the United States and Canada. Volume One, Seventh Edition. Xlibris, Bloomington, Indiana, 613 pp.
    Fowler, H.W. (1906) Note on the Dusky Salamander. Proceedings of the Academy of Natural Sciences of Philadelphia, 58, 356–357.
    Fowler, H.W. & Dunn, E.R. (1917) Notes on Salamanders. Proceedings of the Academy of Natural Sciences of Philadelphia, 69, 7–28.
    Frétey, T. & Raffaëlli, J. (2021) Barton and the forgotten description of Salamandra horrida (Amphibia, Urodela): story of a rediscovery. Bionomina, 25 (1), 35–51. https://doi.org/10.11646/bionomina.25.1.3
    Garden, A. (1821) Correspondence with Linnaeus: Charlestown, June 20, 1771. In: Smith, J.E. (Ed.), A selection of the correspondence of Linnaeus and other naturalists from the original manuscripts. Vol. I. Longman, Hurst, Rees, Orme, and Brown, London, pp. 330–337.
    de Garsault, F.A.P. (1764) Les Figures des Plantes et Animaux d’Usage en Médecine, Décrits dans la Matière Médicale de Mr. Geoffroy Médecin, Dessinés d’après Nature. Niquet scrip. Tome 5. Mrs. Defehrt, Prevost, Duflos, Martinet &c., Paris, 20 pp., pls. 644–729. https://doi.org/10.5962/bhl.title.49481
    Gill, T. (1903a) The first edition of Holbrook’s North American Herpetology. Science, 17, 910–912. https://doi.org/10.1126/science.17.440.910
    Gill, T. (1903b) Biographical memoir of John Edwards Holbrook, 1794–1871. National Academy of Science Memoirs, 1, 49–77.
    Gilliams, J. (1818) Description of two new species of Linnaean Lacerta. Journal of the Academy of Natural Sciences of Philadelphia, 1, 460.
    Gmelin, J.F. (1789) Caroli a Linné, systema naturae. Tom. I. Pars III. Beer, Lipsiae, 483 pp. [pp. 1033–1516]
    Goodwin, G.H. (1960) Unrecorded papers of Rafinesque and Jacob Green. Systematic Zoology, 9 (1), 35–36. https://doi.org/10.2307/2411540
    Gravenhorst, J.L.C. (1807) Vergleichende Uebersicht des Linneischen und einiger neuern zoologischen Systeme ... Nebst dem eingeschalteten Verzeichnisse der zoologischen Sammlung des Verfassers und den Beschreibungen neuer Thierarten, die in derselben vorhanden sind. H. Dieterich, Göttingen, 476 pp.
    Gray, J.E. (1825) A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy, London, Series 2, 10, 193–217.
    Gray, J.E. (1850) Catalogue of the Specimens of Amphibia in the Collection of the British Museum: Part II. Batrachia Gradientia, Etc. Trustees of the British Museum, London, 72 pp.
    Green, J. (1818) Descriptions of several species of North American Amphibia, accompanied with observations. Journal of the Academy of Natural Sciences of Philadelphia, 1, 348–359.
    Green, J. (1825a) Description of a new species of salamander. Journal of the Academy of Natural Sciences of Philadelphia, 5, 116–118.
    Green, J. (1825b) Description of a new species of salamander. The Port Folio, and New York Monthly Magazine, 20, 159.
    Green, J. (1827) An account of some new species of salamanders. Contributions of the Maclurian Lyceum to the Arts and Sciences, 1, 3–8.
    Green, J. (1830) Reclamation of salamanders—in a letter to the Baron F. Cuvier. Transactions of the Albany Institute, 1, 150–151.
    Green, J. (1831) Description of two new species of salamander. Journal of the Academy of Natural Sciences of Philadelphia, 6, 253–255.
    Grobman, A.B. (1950) The distribution of the races of Desmognathus fuscus in the Southern States. Natural History Miscellanea, 70, 1–8.
    Hallowell, E. (1856) Descriptions of two new species of urodeles, from Georgia. Proceedings of the Academy of Natural Sciences of Philadelphia, 8, 130–131.
    Hallowell, E. (1858) On the Caducibranchiate Urodele Batrachians. Proceedings of the Academy of Natural Sciences of Philadelphia, 3, 337–366.
    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, 223–234.
    Harlan, R. (1825b) Further observations on the Amphiuma means. Annals of the Lyceum of Natural History of New-York, 1, 269–270. https://doi.org/10.1111/j.1749-6632.1825.tb00009.x
    Harlan, R. (1825c) Description of a variety of the Coluber fulvius, Linn. a new species of Scincus, and two new species of Salamandra. Journal of the Academy of Natural Sciences of Philadelphia, 5, 154–158.
    Harlan, R. (1825d) Description of a new species of Salamandra. Journal of the Academy of Natural Sciences of Philadelphia, 5, 136.
    Harlan, R. (1826) Notice of a new species of salamander, (inhabiting Pennsylvania.). American Journal of Science and Arts, 10, 286–287.
    Harlan, R. (1827) Genera of North American Reptilia, and synopsis of the species. Journal of the Academy of Natural Sciences of Philadelphia, 5, 317–372.
    Harlan, R. (1829) Description of a new species of Salamandra. Journal of the Academy of Natural Sciences of Philadelphia, 6, 101.
    Harlan, R. (1835) Medical and physical researches, or, Original memoirs in medicine, surgery, physiology, geology, zoology, and comparative anatomy. L.R. Bailey, Philadelphia, Pennsylvania, 653 pp. https://doi.org/10.5962/bhl.title.107564
    Harper, F. (1940) Some works of Bartram, Daudin, Latreille, and Sonnini, and their bearing upon North American herpetological nomenclature. American Midland Naturalist, 23 (3), 692–723. https://doi.org/10.2307/2420453
    Harrison, J.R. & Guttman, S.I. (2003) A new species of Eurycea (Caudata: Plethodontidae) from North and South Carolina. Southeastern Naturalist, 2, 159–178. https://doi.org/10.1656/1528-7092(2003)002[0159:ANSOEC]2.0.CO;2
    Highton, R. (1960) The scientific name of the Red-Backed Salamander. Herpetologica, 16, 236.
    Highton, R. (1961) Erythronota (Salamandra) Rafinesque, 1818; proposed suppression under the plenary powers (Amphibia, Caudata). Bulletin of Zoological Nomenclature, 18, 221–222.
    Highton, R. (1962) Revision of North American salamanders of the Genus Plethodon. Bulletin of the Florida State Museum, 6 (3), 235–367.
    Highton, R., Maha, G.C. & Maxson L.R. (1989) Biochemical evolution in the slimy salamanders of the Plethodon glutinosus complex in the eastern United States. Illinois Biological Monographs, 57, 1–153. https://doi.org/10.5962/bhl.title.49905
    Hime, P.M. (2017) Genomic perspectives on amphibian evolution across multiple phylogenetic scales. PhD Thesis. University of Kentucky, Lexington, Kentucky, 257 pp.
    Holbrook, J.E. (1836) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. I. 1st Edition. J. Dobson, Philadelphia, vi + 120 pp., XXIII pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1838a) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. II. 1st Edition. J. Dobson, Philadelphia, 125 pp., XXVIII pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1838b) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. II. 1st Edition. J. Dobson, Philadelphia, viii + 122 pp., XXX pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1839a) “1836” North American Herpetology: or, A description of the reptiles inhabiting the United States. “Vol. I.” 1st Edition. 2nd Version. J. Dobson, Philadelphia, viii + 132 pp., XXVII pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1839b) “1838” North American Herpetology: or, A description of the reptiles inhabiting the United States. “Vol. II.” 1st Edition. 2nd Version. J. Dobson, Philadelphia, iv + 130 pp., XXX pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1840) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. IV. 1st Edition. J. Dobson, Philadelphia, viii + 126 pp., XXVIII pls. https://doi.org/10.5962/bhl.title.51821
    Holbrook, J.E. (1842a) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. I. 2nd Edition. J. Dobson, Philadelphia, xv + 152 pp., XXIV pls. https://doi.org/10.5962/bhl.title.4997
    Holbrook, J.E. (1842b) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. II. 2nd Edition. J. Dobson, Philadelphia, vi + 142 pp., XX pls. https://doi.org/10.5962/bhl.title.4997
    Holbrook, J.E. (1842c) North American Herpetology: or, A description of the reptiles inhabiting the United States. Second Edition, Volume III. J. Dobson, Philadelphia, ii + 128 pp., XXX pls. https://doi.org/10.5962/bhl.title.4997
    Holbrook, J.E. (1842d) North American Herpetology: or, A description of the reptiles inhabiting the United States. Second Edition, Volume IV. J. Dobson, Philadelphia, Pennsylvania, vi + 138 pp., XXXV pls. https://doi.org/10.5962/bhl.title.4997
    Holbrook, J.E. (1842e) North American Herpetology: or, A description of the reptiles inhabiting the United States. Second Edition, Volume V. J. Dobson, Philadelphia, vi + 118 pp., XXXVIII pls. https://doi.org/10.5962/bhl.title.4997
    Huheey, J.E. & Smith, H.M. (1962) Comments on the proposed suppression of Salamandra erythronota Rafinesque, 1818. Z.N.(S.) 1467. Bulletin of Zoological Nomenclature, 19 (1), 26.
    Huysseune, A., Cerny, R. & Witten, P.E. (2022) The conundrum of pharyngeal teeth origin: the role of germ layers, pouches, and gill slits. Biological Reviews, 97 (1), 414–447. https://doi.org/10.1111/brv.12805
    Jackson, N.D. (2005) Phylogenetic history, morphological parallelism, and speciation in a complex of Appalachian salamanders (Genus: Desmognathus). MS Thesis. Brigham Young University, Provo, Utah, 358 pp.
    Jones, K.S. & Weisrock, D.W. (2018) Genomic data reject the hypothesis of sympatric ecological speciation in a clade of Desmognathus salamanders. Evolution, 72, 2378–2393. https://doi.org/10.1111/evo.13606
    Jones, M.T., Voss, S.R., Ptacek, M.B., Weisrock, D.W. & Tonkyn, D.W. (2006) River drainages and phylogeography: an evolutionary significant lineage of shovel-nosed salamander (Desmognathus marmoratus) in the southern Appalachians. Molecular Phylogenetics and Evolution, 38, 280–287. https://doi.org/10.1016/j.ympev.2005.05.007
    Kaliontzopoulou, A. (2011) Geometric morphometrics in herpetology: modern tools for enhancing the study of morphological variation in amphibians and reptiles. Basic and Applied Herpetology, 25, 5–32. https://doi.org/10.11160/bah.11016
    Kozak, K.H. (2017) What drives variation in plethodontid salamander species richness over space and time? Herpetologica, 73, 220–228. https://doi.org/10.1655/HERPETOLOGICA-D-16-00085.1
    Kozak, K.H., Larson, A., Bonett, R.M. & Harmon, L.J. (2005) Phylogenetic analysis of ecomorphological divergence, community structure, and diversification rates in Dusky Salamanders (Plethodontidae: Desmognathus). Evolution, 59, 2000–2016. https://doi.org/10.1111/j.0014-3820.2005.tb01069.x
    Lacépède, B.G.É. (1788) Histoire Naturelle des Quadrupèdes Ovipares et des Serpens, des Poisson et des Cetaces. 12mo Edition. Vol. II. Hôtel de Thou, Paris, 462 pp. https://doi.org/10.5962/bhl.title.5036
    Lamb, T. & Beamer, D.A. (2012) Digits lost or gained? Evidence for pedal evolution in the Dwarf Salamander complex (Eurycea, Plethodontidae). PLoS ONE, 7, e37544. https://doi.org/10.1371/journal.pone.0037544
    Latreille, P.A. (1800) Histoire naturelle des salamandres de France, précédée d’un tableau méthodique des autres reptiles. Villier, Paris, xlvii + 63 pp., 6 pls. https://doi.org/10.5962/bhl.title.5045
    Laurenti, J.N. (1768) Specimen medicum, exhibens synopsin Reptilium emendatam cum experimentis circa venena et antidota Reptilium austriacorum. Joan Thomae nob. de Trattnern, Viennae, ii + 215 pp., 5 pls. https://doi.org/10.5962/bhl.title.5108
    Le Conte, J. (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, 52–58. https://doi.org/10.1111/j.1749-6632.1824.tb00216.x
    Le Conte, J.E. (1828) Description of a new species of Siren. Annals of the Lyceum of the Natural History of New York, 2, 133–134. https://doi.org/10.1111/j.1749-6632.1826.tb00242.x
    Leuckart, F.S. (1821) Einiges über die fischartigen Amphibien. Isis von Oken, 9, 257–265.
    Linnaeus, C. (1751) Philosophia botanica: in qua explicantur fundamenta botanica cum definitionibus partium, exemplis terminorum, observationibus rariorum, adjectis figuris aeneis. Godofr Kiesewetter, Stockholmiae, 362 pp. https://doi.org/10.5962/bhl.title.37652
    Linnæus, C. (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Salvius, Holmiæ, iv + 824 pp. https://doi.org/10.5962/bhl.title.542
    Linnaeus, C. [Linné, C. a] (1766) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. Salvius, Holmiæ, 532 pp. https://doi.org/10.5962/bhl.title.68927
    Linnaeus, C. [Linné, C. a] (1767) Systema naturæ per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima tertia, ad editionem duodecimam reformatam Holmiensem. Trattnern, Vindobonae, 532 pp. https://doi.org/10.5962/bhl.title.157601
    Malnate, E.V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123, 345–375.
    Means, D.B. (1999) Desmognathus auriculatus. Catalogue of American Amphibians and Reptiles, 681, 1–6.
    Means, D.B., Lamb, J.Y. & Bernardo, J. (2017) A new species of dusky salamander (Amphibia: Plethodontidae: Desmognathus) from the Eastern Gulf Coastal Plain of the United States and a redescription of D. auriculatus. Zootaxa, 4263 (3), 467–506. https://doi.org/10.11646/zootaxa.4263.3.3
    Mittleman, M.B. (1947) American Caudata. I. Geographic Variation in Manculus quadridigitatus. Herpetologica, 3, 209–224.
    Mittleman, M.B. (1951) American Caudata. VII. Two new salamanders of the genus Plethodon. Herpetologica, 7, 105–112.
    Mittleman, M.B. (1966) Eurycea bislineata. Catalogue of American Amphibians and Reptiles, 45, 1–4.
    Mittleman, M.B. (1967) Manculus and M. quadridigitatus. Catalogue of American Amphibians and Reptiles, 44, 1–2.
    Moore, J.P. (1899) Leurognathus marmorata, a new genus and species of salamander of the family Desmognathidæ. Proceedings of the Academy of Natural Sciences of Philadelphia, 51, 316–323.
    O’Connell, K.A., Mulder, K.P., Wynn, A., de Queiroz, K. & Bell, R.C. (2022) Genomic library preparation and hybridization capture of formalin-fixed tissues and allozyme supernatant for population genomics and considerations for combining capture- and RADseq-based single nucleotide polymorphism data sets. Molecular Ecology Resources, 22, 487–502. https://doi.org/10.1111/1755-0998.13481
    Osgood, W.H. (1928) Why is a type specimen. Journal of Mammalogy, 9, 52–56. https://doi.org/10.2307/1373357
    Österdam, A. (1766) Siren lacertina. Dissertatione academica, Upsaliae, 20 pp., 1 pl.
    Petranka, J.W. (2010) Salamanders of the United States and Canada. Smithsonian Books, Washington, 592 pp.
    Phillips, V.T. & Phillips M.E. (1963) Guide to the manuscript collections in the Academy of Natural Sciences of Philadelphia. Edwards, Ann Arbor, Michigan, 553 pp.
    Powell, R., Conant, R. & Collins, J.T. (2016) Peterson field guide to reptiles and amphibians of eastern and central North America. Houghton Mifflin Harcourt, Boston, 512 pp.
    Pyron, R.A. & Beamer, D.A. (2020) The herpetological legacy of Jacob Green and the nomenclature of some North American lizards and salamanders. Zootaxa, 4838 (2), 221–247. https://doi.org/10.11646/zootaxa.4838.2.4
    Pyron, R.A., O’Connell, K.A., Lemmon, E.M., Lemmon, A.R. & Beamer, D.A. (2020) Phylogenomic data reveal reticulation and incongruence among mitochondrial candidate species in Dusky Salamanders (Desmognathus). Molecular Phylogenetics and Evolution, 146, 106751. https://doi.org/10.1016/j.ympev.2020.106751
    Pyron, R.A., Gotte, S. & Burbrink, F.T. (2021) Additional comments on the types and nomina of several North American ratsnakes (Pantherophis obsoletus complex, Colubridae, Serpentes). Bionomina, 25 (1), 70–75. https://doi.org/10.11646/bionomina.25.1.5
    Pyron, R.A., Beamer, D.A., Holzheuser, C.R., Lemmon, E.M., Lemmon, A.R., Wynn, A.H. & O’Connell, K.A. (2022a) Contextualizing enigmatic extinctions using genomic DNA from fluid-preserved museum specimens of Desmognathus salamanders. Conservation Genetics, 23, 375–386. https://doi.org/10.1007/s10592-021-01424-4
    Pyron, R.A., O’Connell, K.A., Lemmon, E.M., Lemmon, A.R. & Beamer, D.A. (2022b) Candidate-species delimitation in Desmognathus salamanders reveals gene flow across lineage boundaries, confounding phylogenetic estimation and clarifying hybrid zones. Ecology and Evolution, 12 (2), e8574. https://doi.org/10.1002/ece3.8574
    Rafinesque, C.S. (1815) Analyse de Nature, ou Tableau de l’Universe et des Corps Organisés. Jean Barravecchia, Palermo, 244 pp. https://doi.org/10.5962/bhl.title.106607
    Rafinesque, C.S. (1818a) Description of a new American salamander—the red-backed salamander from the Highlands. Scientific Journal of New York, 1, 25–26.
    Rafinesque, C.S. (1818b) Farther accounts of discoveries in natural history, in the western states. American Monthly Magazine and Critical Review, 4, 39–42.
    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.
    Rafinesque C.S. (1820) III Class. Erpetia.–The reptiles. Annals of Nature, 1, 4–6. https://doi.org/10.5962/bhl.title.106763
    Rafinesque, C.S. (1822) On two new salamanders of Kentucky. Kentucky Gazette, New Series, 1 (9), 3.
    Reed, C.F. (1960) Plethodon erythronotus (Raf.), the Red-Backed Salamander. Herpetologica, 16, 207–213.
    Rossman, D.A. (1959) Ecosystematic Relationships of the Salamanders Desmognathus fuscus auriculatus Holbrook and Desmognathus fuscus carri Neill. Herpetologica, 15, 149–155.
    Roux-Estéve, R. (1976) Liste des spécimens donnés par Holbrook encore en collection au Muséum national d’Histoire naturelle de Paris. Private papers (unpublished manuscript), Muséum national d’Histoire naturelle, Paris, 12 pp.
    Sanders, A.E. & Anderson, W.D. (1999) Natural history investigations in South Carolina: from colonial times to the present. University of South Carolina Press, Columbia, 333 pp.
    Say, T. (1819) Notes on herpetology. The American Journal of Science, &c, 1 (2), 256–265.
    Say, T. (1823) Coluber obsoletus. In: James, E. (Ed.), “1822” Account of an expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and ‘20, by order of the Hon. J. C. Calhoun, Sec’y of War: under the command of Major Stephen H. Long. From the notes of Major Long, Mr. T. Say, and other gentlemen of the exploring party. Compiled by Edwin James, botanist and geologist for the expedition. In two vols. – with an atlas. H.C. Carey and I. Lea, Philadelphia, pp. 140–141.
    Schmidt, K.P. (1953) A Check List of North American Amphibians and Reptiles. 6th Edition. University of Chicago Press, Chicago, 280 pp.
    Schneider, J.G. (1799) Historiae Amphibiorum Naturalis et Literarariae. Fasciculus Primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species Descriptos Notisque suis Distinctos. Friederici Frommanni, Jena, 264 pp., 2 pls.
    Seiss, C.F. (1881) The Spotted Amblystome. Scientific American, 45, 329. https://doi.org/10.1038/scientificamerican11191881-329
    Shaw, G. (1802) General Zoology or Systematic Natural History. Vol. III. Amphibia. G. Kearsley, London, 615 pp.
    Shimek, B. (1924) Holbrook’s North American Herpetology. Proceedings of the Iowa Academy of Science, 31, 427–430.
    Simmons, J.E. (2014) Fluid preservation: a comprehensive reference. Rowman & Littlefield, Lanham, Maryland, 347 pp.
    Smith, H.M., Smith, R.B. & Sawin, H.L. (1975) The authorship and date of publication of Siren intermedia (Amphibia: Caudata). The Great Basin Naturalist, 35, 100–102.
    Smith P.W. (1963) Plethodon cinereus. Catalogue of American Amphibians and Reptiles, 5, 1–3.
    Sonnini de Manoncourt, C.S. & Latreille, P.A. (1801a) Histoire naturelle des Reptiles. Tome 2. Première partie. Quadrupèdes et Bipèdes Ovipares. Deterville, Paris, iii + 332 pp., 19 pls.
    Sonnini de Manoncourt, C.S. & Latreille, P.A. (1801b) Histoire naturelle des Reptiles. Tome 4. Seconde partie. Serpens. Deterville, Paris, iii + 410 pp., 14 pls.
    Stejneger, L. (1902) A salamander new to the District of Columbia. Proceedings of the Biological Society of Washington, 15, 239–240.
    Stejneger, L. (1903) Rediscovery of one of Holbrook’s salamanders. Proceedings of the United States National Museum, 26, 557–558. https://doi.org/10.5479/si.00963801.1321.557
    Stephens, L.D. (1975) Of Mercury, Moses, and Medicine: Views of Dr. John M. B. Harden. The Georgia Historical Quarterly, 59, 402–415.
    Stephens, L.D. (1997) John Edwards Holbrook (1794–1871) and Lewis Reeve Gibbes (1810–1894): exemplary naturalists in the Old South. In: Pietsch, T.W. & Anderson, W.D. (Eds.), Collection Building in Ichthyology and Herpetology, Special Publication No. 3. American Society of Ichthyologists and Herpetologists, Lawrence, Kansas, pp. 447–458.
    Stephens, L.D. (2000) Science, race, and religion in the American South: John Bachman and the Charleston circle of naturalists, 1815–1895. University of North Carolina Press, Chapel Hill, North Carolina, 360 pp.
    Storer, D.H. (1840) A report on the reptiles of Massachusetts. Boston Journal of Natural History, 3, 1–64.
    Temminck, C.J. & Schlegel, H. (1838) Fauna Japonica sive Descriptio animalium, quae in itinere per Japonianum, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenent, suscepto, annis 1823–1830 colleget, notis observationibus et adumbrationibus illustratis. Vol. 3. Chelonia, Ophidia, Sauria, Batrachia. J.G. Lalau, Leiden, 144 pp.
    Thireau, M. (1986) Catalogue des types d’urodeÌles du Museìum national d’histoire naturelle: revue critique. Museum National d’Histoire Naturelle, Paris, 92 pp.
    Tschudi, J.J. (1838) Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Petitpierre, Neuchâtel, ii + 98 pp., 6 pls. https://doi.org/10.5962/bhl.title.4883
    Uzzell, T. (1967) Ambystoma jeffersonianum. Catalogue of American Amphibians and Reptiles, 47, 1–2.
    Valentine, B.D. (1963) The salamander genus Desmognathus in Mississippi. Copeia, 1963, 130–139. https://doi.org/10.2307/1441280
    Valentine, B.D. (1974) Desmognathus quadramaculatus. Catalogue of American Amphibians and Reptiles, 153, 1–4.
    Waite, F.C. (1907) Specific name of Necturus maculosus. The American Naturalist, 41 (481), 23–30. https://doi.org/10.1086/278699
    Wooten, J.A. (2008) Comparative phylogeography of dusky salamanders, genus Desmognathus, in the southern Appalachian Mountains. PhD Thesis. University of Alabama, Tuscaloosa, Alabama, 192 pp.
    Wooten, J.A. & Rissler, L.J. (2011) Ecological associations and genetic divergence in Black-bellied Salamanders (Desmognathus quadramaculatus) of the Southern Appalachian Mountains. Acta Herpetologica, 6, 175–208.
    Worthington, R.D. (1976) Current nomenclature for scientific names in Holbrook’s “North American Herpetology” including a Table of Contents. In: Adler, K.A. (Ed.), Holbrook’s North American Herpetology by John Edwards Holbrook. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, pp. xlv–xlix.
    Worthington, R.D. & Worthington, P.H. (1976) John Edwards Holbrook, father of American herpetology. In: Adler, K.A. (Ed.), Holbrook’s North American Herpetology by John Edwards Holbrook. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, pp. xiii–xxvii.
    Wray, K.P., Means, D.B. & Steppan, S.J. (2017) Revision of the Eurycea quadridigitata (Holbrook 1842) complex of Dwarf Salamanders (Caudata: Plethodontidae: Hemidactyliinae) with a description of two new species. Herpetological Monographs, 31, 18–46. https://doi.org/10.1655/HERPMONOGRAPHS-D-16-00011
    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, 1–249. https://doi.org/10.5479/si.03629236.24.1