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<document id="1C205CD9A8E80113377EA12BBBEA7143" ID-DOI="10.11646/zootaxa.5134.2.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="6534805" ID-ZooBank="3C3F497E-7B50-4E49-8983-D773581F18FD" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="carolina" checkinTime="1652171733176" checkinUser="plazi" docAuthor="Pyron, R. Alexander &amp; Beamer, David A." docDate="2022" docId="DF5187BB533FFFE3FF588B38FDCAD045" docLanguage="en" docName="zootaxa.5134.2.1.pdf" docOrigin="Zootaxa 5134 (2)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Salamandra Garsault 1764" docType="treatment" docVersion="1" lastPageNumber="174" masterDocId="2368FFC35329FFF4FFCF8D6DFFA9D37F" masterDocTitle="A nomenclatural and taxonomic review of the salamanders (Urodela) from Holbrooks North American Herpetology" masterLastPageNumber="196" masterPageNumber="151" pageNumber="173" updateTime="1734705238574" updateUser="carolina" zenodo-license-document="CLOSED">
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<mods:title id="3DA85B19530F85705B642746AD11C06B">A nomenclatural and taxonomic review of the salamanders (Urodela) from Holbrooks North American Herpetology</mods:title>
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<mods:affiliation id="CA7CE00BD599F2B2D50A8EA885B08896">Department of Biological Sciences, The George Washington University, 2023 G St. NW, Washington, DC 20052 rpyron @ colubroid. org; https: // orcid. org / 0000 - 0003 - 2524 - 1794 &amp; Division of Amphibians and Reptiles, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560</mods:affiliation>
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<taxonomicName id="90F84D2E533FFFE2FF588B38FE89D510" ID-CoL="7BSB" authority="Haldemani" authorityName="Garsault" authorityYear="1764" box="[151,288,1621,1647]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="22" pageNumber="173" phylum="Chordata" rank="genus">Salamandra</taxonomicName>
“Haldemani”
</emphasis>
<bibRefCitation id="33694B5C533FFFE2FE058B38FD06D50F" author="Holbrook, J. E." box="[458,687,1621,1648]" pageId="22" pageNumber="173" refId="ref32433" refString="Holbrook, J. E. (1840) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. IV. 1 st Edition. J. Dobson, Philadelphia, viii + 126 pp., XXVIII pls. https: // doi. org / 10.5962 / bhl. title. 51821" type="book" year="1840">Holbrook, 1840:125</bibRefCitation>
, pl. 28
</paragraph>
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<paragraph id="574736AD533FFFE2FCCD8B3BFC60D50F" blockId="22.[151,1437,1621,2008]" box="[770,969,1621,1648]" pageId="22" pageNumber="173">
[=
<emphasis id="658CEABF533FFFE2FCD48B3BFC69D510" box="[795,960,1621,1647]" italics="true" pageId="22" pageNumber="173">nomen dubium</emphasis>
]
</paragraph>
<paragraph id="574736AD533FFFE2FF088B17FD87D4CC" blockId="22.[151,1437,1621,2008]" pageId="22" pageNumber="173">
<collectorName id="FA0D537B533FFFE2FF088B17FF47D5EB" box="[199,238,1658,1684]" pageId="22" pageNumber="173">For</collectorName>
the final salamander and last account of the initially overlooked fourth volume of the first edition,
<bibRefCitation id="33694B5C533FFFE2FAFE8B14FF4BD5C8" author="Holbrook, J. E." pageId="22" pageNumber="173" refId="ref32433" refString="Holbrook, J. E. (1840) North American Herpetology: or, A description of the reptiles inhabiting the United States. Vol. IV. 1 st Edition. J. Dobson, Philadelphia, viii + 126 pp., XXVIII pls. https: // doi. org / 10.5962 / bhl. title. 51821" type="book" year="1840">Holbrook (1840)</bibRefCitation>
gave an ambiguous description of a new species based on specimens he had received from
<collectingRegion id="953CF84F533FFFE2FACF8BF0FEAED5A4" isEnumeration="true" pageId="22" pageNumber="173">Pennsylvania, Maryland</collectingRegion>
, and
<collectingRegion id="953CF84F533FFFE2FE8A8BACFE0DD5A4" box="[325,420,1729,1755]" country="United States of America" name="Virginia" pageId="22" pageNumber="173">Virginia</collectingRegion>
.
<collectorName id="FA0D537B533FFFE2FE7E8BACFE49D5A4" box="[433,480,1729,1755]" pageId="22" pageNumber="173">One</collectorName>
specimen in particular came from
<collectorName id="FA0D537B533FFFE2FCBC8BACFB81D5A4" box="[883,1064,1729,1755]" pageId="22" pageNumber="173">S. S. Haldeman</collectorName>
, collected from “the borders” of the
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in
<collectingRegion id="953CF84F533FFFE2FE708B88FDFED580" box="[447,599,1765,1791]" country="United States of America" name="Pennsylvania" pageId="22" pageNumber="173">Pennsylvania</collectingRegion>
, for whom
<collectorName id="FA0D537B533FFFE2FD178B88FCEAD580" box="[728,835,1765,1791]" pageId="22" pageNumber="173">Holbrook</collectorName>
named the species.
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are thus numerous primary
<typeStatus id="8843880F533FFFE2FF588A67FF55D45B" box="[151,252,1802,1828]" pageId="22" pageNumber="173" type="syntype">syntypes</typeStatus>
, one of which was illustrated by
<collectorName id="FA0D537B533FFFE2FDA98A64FCD4D45B" box="[614,893,1801,1828]" pageId="22" pageNumber="173">Thomas M. Logan, M.D.</collectorName>
in modest detail. The description and illustration were reproduced essentially verbatim by
<bibRefCitation id="33694B5C533FFFE2FDA38A40FCFDD437" author="Holbrook, J. E." box="[620,852,1837,1864]" pageId="22" pageNumber="173" refId="ref32745" refString="Holbrook, J. E. (1842 e) 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" type="book" year="1842">Holbrook (1842e:58</bibRefCitation>
, pl. 18).
<bibRefCitation id="33694B5C533FFFE2FC748A40FBC2D437" author="Schmidt, K. P." box="[955,1131,1837,1864]" pageId="22" pageNumber="173" refId="ref35329" refString="Schmidt, K. P. (1953) A Check List of North American Amphibians and Reptiles. 6 th Edition. University of Chicago Press, Chicago, 280 pp." type="book" year="1953">Schmidt (1953)</bibRefCitation>
restricted the type locality to Harrisburg,
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, without further justification. None of the type series was ever noted to exist by later authors or encountered by us;
<bibRefCitation id="33694B5C533FFFE2FE248A18FD1BD4EF" author="Mittleman, M. B." box="[491,690,1909,1936]" pageId="22" pageNumber="173" pagination="1 - 4" refId="ref34085" refString="Mittleman, M. B. (1966) Eurycea bislineata. Catalogue of American Amphibians and Reptiles, 45, 1 - 4." type="journal article" year="1966">Mittleman (1966)</bibRefCitation>
stated that the specimen figured may be fixed as the
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, but Holbrook gave no indication of this.
</paragraph>
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This name is yet a
<emphasis id="658CEABF533FFFE2FE5D8AD3FD96D4A8" box="[402,575,1981,2007]" italics="true" pageId="22" pageNumber="173">nomen dubium;</emphasis>
<bibRefCitation id="33694B5C533FFFE2FD8B8AD0FD66D4A8" author="Cope, E. D." box="[580,719,1981,2007]" pageId="22" pageNumber="173" pagination="1 - 525" refId="ref29724" refString="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" type="journal article" year="1889">Cope (1889)</bibRefCitation>
reported that Baird considered it to be a synonym of
<taxonomicName id="90F84D2E533FFFE3FADC8AD0FE34D3CE" authority="Green, 1818" authorityName="Green" authorityYear="1818" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" lastPageId="23" lastPageNumber="174" order="Caudata" pageId="22" pageNumber="173" phylum="Chordata" rank="species" species="bislineata">
<emphasis id="658CEABF533FFFE3FADC8AD0FEACD3CE" italics="true" lastPageId="23" lastPageNumber="174" pageId="22" pageNumber="173">Salamandra bislineata</emphasis>
<bibRefCitation id="33694B5C533EFFE3FEC18DFAFE34D3CE" author="Green, J." box="[270,413,151,177]" pageId="23" pageNumber="174" pagination="348 - 359" refId="ref31172" refString="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." type="journal article" year="1818">Green, 1818</bibRefCitation>
</taxonomicName>
. This was repeated by
<bibRefCitation id="33694B5C533EFFE3FD6D8DFAFC9CD3CD" author="Dunn, E. R." box="[674,821,151,178]" pageId="23" pageNumber="174" refId="ref30281" refString="Dunn, E. R. (1926) The salamanders of the family Plethodontidae. Smith College, Northampton, Massachusetts, 446 pp." type="book" year="1926">Dunn (1926)</bibRefCitation>
, who stated that it was based on a transforming larva, which was accepted by
<bibRefCitation id="33694B5C533EFFE3FE648DD6FDDFD3A9" author="Mittleman, M. B." box="[427,630,187,214]" pageId="23" pageNumber="174" pagination="1 - 4" refId="ref34085" refString="Mittleman, M. B. (1966) Eurycea bislineata. Catalogue of American Amphibians and Reptiles, 45, 1 - 4." type="journal article" year="1966">Mittleman (1966)</bibRefCitation>
. However,
<bibRefCitation id="33694B5C533EFFE3FD358DD6FC34D3A9" author="Fowler, H. W." box="[762,925,187,214]" pageId="23" pageNumber="174" pagination="356 - 357" refId="ref30591" refString="Fowler, H. W. (1906) Note on the Dusky Salamander. Proceedings of the Academy of Natural Sciences of Philadelphia, 58, 356 - 357." type="journal article" year="1906">Fowler (1906)</bibRefCitation>
notes that the description of the pattern and length of the tail do not match the Northern Two-lined Salamander; furthermore, the total length is given as
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, much larger than metamorphosing
<taxonomicName id="90F84D2E533EFFE3FD4B8C6EFD48D263" authorityName="Rafinesque" authorityYear="1822" box="[644,737,259,284]" class="Amphibia" family="Plethodontidae" genus="Eurycea" kingdom="Animalia" order="Caudata" pageId="23" pageNumber="174" phylum="Chordata" rank="genus">
<emphasis id="658CEABF533EFFE3FD4B8C6EFD48D263" box="[644,737,259,284]" italics="true" pageId="23" pageNumber="174">Eurycea</emphasis>
</taxonomicName>
. Instead, Fowler notes (and we agree) that the color pattern and size match
<taxonomicName id="90F84D2E533EFFE3FE878C4AFD9DD23E" authority="Green, 1818" authorityName="Green" authorityYear="1818" box="[328,564,295,321]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="23" pageNumber="174" phylum="Chordata" rank="species" species="fusca">
<emphasis id="658CEABF533EFFE3FE878C4AFE09D23E" box="[328,416,295,321]" italics="true" pageId="23" pageNumber="174">S. fusca</emphasis>
<bibRefCitation id="33694B5C533EFFE3FE668C4AFD9DD23E" author="Green, J." box="[425,564,295,321]" pageId="23" pageNumber="174" pagination="348 - 359" refId="ref31172" refString="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." type="journal article" year="1818">Green, 1818</bibRefCitation>
</taxonomicName>
more closely. Therefore, it is more likely that this taxon represents Northern Dusky Salamanders than Northern Two-lined Salamanders, or perhaps a mixture of the two, possibly including metamorphosing larvae of one or both species.
</paragraph>
<paragraph id="574736AD533EFFE3FF088CFEFAF6D166" blockId="23.[151,1436,151,538]" pageId="23" pageNumber="174">
While the first edition was popular and well-received, Holbrook remained dissatisfied with its presentation and organization, and immediately commenced a second edition with many new illustrations (see
<bibRefCitation id="33694B5C533EFFE3FB2F8CDAFECFD289" author="Worthington, R. D. &amp; Worthington, P. H." pageId="23" pageNumber="174" pagination="pp. xiii - xxvii" refId="ref36387" refString="Worthington, R. D. &amp; 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." type="book chapter" year="1976">Worthington and Worthington 1976</bibRefCitation>
). In it (
<bibRefCitation id="33694B5C533EFFE3FE7E8CB6FDC4D28A" author="Holbrook, J. E." box="[433,621,475,501]" pageId="23" pageNumber="174" refId="ref32745" refString="Holbrook, J. E. (1842 e) 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" type="book" year="1842">Holbrook 1842e</bibRefCitation>
), he reprinted all of the accounts from the first edition (often with changes or additions as noted above), and added 14 additional accounts and descriptions, two of them new. These are:
</paragraph>
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<emphasis id="658CEABF533EFFE3FF588F2AFE76D11E" box="[151,479,583,609]" italics="true" pageId="23" pageNumber="174">
<taxonomicName id="90F84D2E533EFFE3FF588F2AFE89D11E" authority="Jeffersoniana" authorityName="Garsault" authorityYear="1764" box="[151,288,583,609]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="23" pageNumber="174" phylum="Chordata" rank="genus">Salamandra</taxonomicName>
<taxonomicName id="90F84D2E533EFFE3FEFE8F2AFE7CD11E" authority="Green, 1827" authorityName="Green" authorityYear="1827" box="[305,469,583,609]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="23" pageNumber="174" phylum="Chordata" rank="species" species="jeffersoniana">Jeffersoniana</taxonomicName>
</emphasis>
<bibRefCitation id="33694B5C533EFFE3FE298F2AFDD9D11E" author="Green, J." box="[486,624,583,609]" pageId="23" pageNumber="174" pagination="3 - 8" refId="ref31270" refString="Green, J. (1827) An account of some new species of salamanders. Contributions of the Maclurian Lyceum to the Arts and Sciences, 1, 3 - 8." type="journal article" year="1827">Green, 1827</bibRefCitation>
</paragraph>
<paragraph id="574736AD533EFFE3FF088F06FDCAD045" blockId="23.[151,1437,583,826]" pageId="23" pageNumber="174">
<bibRefCitation id="33694B5C533EFFE3FF088F06FE19D1FA" author="Holbrook, J. E." box="[199,432,618,645]" pageId="23" pageNumber="174" refId="ref32745" refString="Holbrook, J. E. (1842 e) 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" type="book" year="1842">Holbrook (1842e:51</bibRefCitation>
, pl. 14) gave a brief account of the Jefferson Salamander (
<taxonomicName id="90F84D2E533EFFE3FB978F06FA25D1FA" baseAuthorityName="Green" baseAuthorityYear="1827" box="[1112,1420,619,645]" class="Amphibia" family="Ambystomatidae" genus="Ambystoma" kingdom="Animalia" order="Caudata" pageId="23" pageNumber="174" phylum="Chordata" rank="species" species="jeffersonianum">
<emphasis id="658CEABF533EFFE3FB978F06FA25D1FA" box="[1112,1420,619,645]" italics="true" pageId="23" pageNumber="174">Ambystoma jeffersonianum</emphasis>
</taxonomicName>
), which he stated he had never seen. He also stated that the rough illustration by J. Queen was copied from T. R. Peales original drawing of the
<typeStatus id="8843880F533EFFE3FE308FDEFDC8D1B2" box="[511,609,691,717]" pageId="23" pageNumber="174" type="holotype">holotype</typeStatus>
figured in
<bibRefCitation id="33694B5C533EFFE3FD2E8FDEFCD5D1B2" author="Green, J." box="[737,892,691,717]" pageId="23" pageNumber="174" pagination="3 - 8" refId="ref31270" refString="Green, J. (1827) An account of some new species of salamanders. Contributions of the Maclurian Lyceum to the Arts and Sciences, 1, 3 - 8." type="journal article" year="1827">Green (1827)</bibRefCitation>
. Recent evidence suggests that this specimen is extant, USNM 3968 (see
<bibRefCitation id="33694B5C533EFFE3FE7E8FBAFD68D18E" author="Pyron, R. A. &amp; Beamer, D. A." box="[433,705,727,753]" pageId="23" pageNumber="174" pagination="221 - 247" refId="ref34425" refString="Pyron, R. A. &amp; 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" type="journal article" year="2020">Pyron and Beamer 2020</bibRefCitation>
: figs. 9, 10). Thus, Holbrook figured a specimen that is still extant but which he never saw, a condition that also potentially holds for USNM 3968, USNM 3981, USNM 3840, and MNHN-RA0.7871 (see accounts below).
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