<document id="D92B5A026A4D1DC1DE429E6D8C8346A9" ID-DOI="10.1111/j.1096-3642.2007.00244.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5428560" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630417322133" checkinUser="felipe" docAuthor="Mutter, Raoul J., Blanger, Keith De &amp; Neuman, Andrew G." docDate="2007" docId="692EA02D8F5DFFC4FE8AFE528A15DAEA" docLanguage="en" docName="j.1096-3642.2007.00244.x.pdf" docOrigin="Zoological Journal of the Linnean Society 149 (3)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2007.00244.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Palaeobates : PIMUZ T 1179" docType="treatment" docVersion="4" lastPageNumber="335" masterDocId="9517D8558F44FFDEFF9CFFC2884ED971" masterDocTitle="Elasmobranchs from the Lower Triassic Sulphur Mountain Formation near Wapiti Lake (BC, Canada)" masterLastPageNumber="337" masterPageNumber="309" pageNumber="334" updateTime="1705831329446" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title id="BC747BDFE84FB8D3372BDB521A3EE296">Elasmobranchs from the Lower Triassic Sulphur Mountain Formation near Wapiti Lake (BC, Canada)</mods:title>
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<mods:namePart id="BE3768CD8DABF9E10170D783B5A8E46A">Mutter, Raoul J.</mods:namePart>
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<paragraph id="E138113B8F5DFFC7FE8AFE528AD8D8D6" blockId="25.[278,662,400,424]" box="[278,662,400,424]" pageId="25" pageNumber="334">
<heading id="BA70A6578F5DFFC7FE8AFE528AD8D8D6" box="[278,662,400,424]" centered="true" fontSize="9" level="2" pageId="25" pageNumber="334" reason="2">
COMPARISON WITH 
<taxonomicName id="26876AB88F5DFFC7FE6AFE538AD8D8D6" authorityName=": PIMUZ T" authorityYear="1179" box="[502,662,401,424]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FE6AFE538AD8D8D6" box="[502,662,401,424]" italics="true" pageId="25" pageNumber="334">PALAEOBATES</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<paragraph id="E138113B8F5DFFC7FF3EFE7B89ABDBB5" blockId="25.[162,779,441,1904]" pageId="25" pageNumber="334">
Several specimens from the Vega-Phroso Siltstone Member of Wapiti Lake consist of fragmentary sections of the torsi and/or dorsal fins with neither teeth nor cranial morphology preserved. One of these very incomplete specimens, CMN 9980, consists mainly of patches of denticles and is preserved in outline only. Lacking most fins and all teeth, it was referred to 
<taxonomicName id="26876AB88F5DFFC7FF3EFD4D8948DBD5" baseAuthorityName="Agassiz" baseAuthorityYear="1837" box="[162,262,655,676]" class="Chondrichthyes" family="Hybodontidae" genus="Hybodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FF3EFD4D8948DBD5" box="[162,262,655,676]" italics="true" pageId="25" pageNumber="334">Hybodus</emphasis>
</taxonomicName>
by 
<bibRefCitation id="85166CCA8F5DFFC7FEADFD4D89A7DBD4" author="Gardiner BG" box="[305,489,655,677]" pageId="25" pageNumber="334" pagination="1 - 154" refId="ref19476" refString="Gardiner BG. 1966. Catalogue of Canadian fossil fishes. Royal Ontario Museum, University of Toronto. Life Sciences Contributions 68: 1 - 154." type="journal article" year="1966">Gardiner (1966)</bibRefCitation>
and to cf. 
<taxonomicName id="26876AB88F5DFFC7FDFDFD4D8AA8DBD5" authorityName=": PIMUZ T" authorityYear="1179" box="[609,742,655,676]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FDFDFD4D8AA8DBD5" box="[609,742,655,676]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
by 
<bibRefCitation id="85166CCA8F5DFFC7FF3EFD6C89AFDBB5" author="Schaeffer B &amp; Mangus M" box="[162,481,686,708]" pageId="25" pageNumber="334" pagination="519 - 563" refId="ref20666" refString="Schaeffer B, Mangus M. 1976. An Early Triassic fish assemblage from British Columbia. Bulletin of the American Museum of Natural History 156: 519 - 563." type="journal article" year="1976">Schaeffer &amp; Mangus (1976)</bibRefCitation>
.
</paragraph>
<paragraph id="E138113B8F5DFFC7FF27FD0F89D4DD64" blockId="25.[162,779,441,1904]" pageId="25" pageNumber="334">
Comparison between 
<taxonomicName id="26876AB88F5DFFC7FE56FD0F8A04DB93" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[458,586,717,738]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="D3F3CD298F5DFFC7FE56FD0F8A04DB93" box="[458,586,717,738]" italics="true" pageId="25" pageNumber="334">Wapitiodus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FDC2FD0F8A83DB93" box="[606,717,717,738]" pageId="25" pageNumber="334" rank="genus">gen. nov.</taxonomicNameLabel>
and 
<taxonomicName id="26876AB88F5DFFC7FF3EFD29891ADA70" box="[162,340,747,769]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="D3F3CD298F5DFFC7FF3EFD298969DA71" box="[162,295,747,768]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
sp.
</taxonomicName>
, the only previously mentioned hybodont taxon from the Sulphur Mountain Formation, stands on weak grounds as the original material and description are insufficient. 
<taxonomicName id="26876AB88F5DFFC7FE7DFC858A28DA2D" authorityName=": PIMUZ T" authorityYear="1179" box="[481,614,839,860]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FE7DFC858A28DA2D" box="[481,614,839,860]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
is now known by various skeletal elements and differs from the material described here in many features in hybodont fin spine structure and in tooth structure (discussed below). 
<taxonomicName id="26876AB88F5DFFC7FF66FC0089CDDAA6" baseAuthorityName="Jaekel" baseAuthorityYear="1889" box="[250,387,962,983]" class="Chondrichthyes" family="Hybodontidae" genus="Polyacrodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FF66FC0089CDDAA6" box="[250,387,962,983]" italics="true" pageId="25" pageNumber="334">Polyacrodus</emphasis>
</taxonomicName>
, 
<taxonomicName id="26876AB88F5DFFC7FE0DFC008A58DAA6" authorityName=": PIMUZ T" authorityYear="1179" box="[401,534,962,983]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FE0DFC008A58DAA6" box="[401,534,962,983]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
and 
<taxonomicName id="26876AB88F5DFFC7FDCEFC008A9CDAA6" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[594,722,962,983]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="D3F3CD298F5DFFC7FDCEFC008A9CDAA6" box="[594,722,962,983]" italics="true" pageId="25" pageNumber="334">Wapitiodus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FD46FC008881DA87" pageId="25" pageNumber="334" rank="genus">gen. nov.</taxonomicNameLabel>
share one plesiomorphic feature, the thick singlecrystallite enameloid.
</paragraph>
<paragraph id="E138113B8F5DFFC7FF27FBDC8AC8DC58" blockId="25.[162,779,441,1904]" pageId="25" pageNumber="334">
<bibRefCitation id="85166CCA8F5DFFC7FF27FBDC89B6DD45" author="Schaeffer B &amp; Mangus M" box="[187,504,1054,1076]" pageId="25" pageNumber="334" pagination="519 - 563" refId="ref20666" refString="Schaeffer B, Mangus M. 1976. An Early Triassic fish assemblage from British Columbia. Bulletin of the American Museum of Natural History 156: 519 - 563." type="journal article" year="1976">Schaeffer &amp; Mangus (1976)</bibRefCitation>
described four (or multiple)-pronged acuminate projections in the dermal denticles that are possibly conspecific with ‘genus A’ (Superorder 
<emphasis id="D3F3CD298F5DFFC7FEADFBB98987DDFE" box="[305,457,1146,1168]" italics="true" pageId="25" pageNumber="334">incertae sedis</emphasis>
) described here. In the presence of conspicuous diverging prongs, the denticles resemble the shape of the crown in certain denticles of 
<taxonomicName id="26876AB88F5DFFC7FF3EFB148ACEDD9D" authority="(Dick, 1998)" baseAuthorityName="Dick" baseAuthorityYear="1998" box="[162,640,1238,1260]" class="Chondrichthyes" family="Sphenacanthidae" genus="Sphenacanthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ctenacanthiformes" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="serrulatus">
<emphasis id="D3F3CD298F5DFFC7FF3EFB148996DD9A" box="[162,472,1238,1259]" italics="true" pageId="25" pageNumber="334">Sphenacanthus serrulatus</emphasis>
(
<bibRefCitation id="85166CCA8F5DFFC7FE6EFB148A37DD9D" author="Dick JRF" box="[498,633,1238,1260]" pageId="25" pageNumber="334" pagination="9 - 25" refId="ref19325" refString="Dick JRF. 1998. Sphenacanthus, a Palaeozoic freshwater shark. Zoological Journal of the Linnean Society 122: 9 - 25." type="journal article" year="1998">Dick, 1998</bibRefCitation>
)
</taxonomicName>
. The welldeveloped multiforaminate pedicles, however, clearly distinguished ‘genus A’ from 
<taxonomicName id="26876AB88F5DFFC7FE72FAD68A30DC59" baseAuthorityName="Dick" baseAuthorityYear="1998" box="[494,638,1299,1321]" class="Chondrichthyes" family="Sphenacanthidae" genus="Sphenacanthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ctenacanthiformes" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="serrulatus">
<emphasis id="D3F3CD298F5DFFC7FE72FAD68A30DC59" box="[494,638,1299,1321]" italics="true" pageId="25" pageNumber="334">S. serrulatus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="E138113B8F5DFFC7FF27FAF08985DF84" blockId="25.[162,779,441,1904]" pageId="25" pageNumber="334">
A similar 
<typeStatus id="3E3CAF998F5DFFC7FEB1FAF18910DC39" box="[301,350,1331,1352]" pageId="25" pageNumber="334">type</typeStatus>
of platform with several long diverging ridges and a well-developed pedicle has, to our knowledge, only been described under the nominal name 
<taxonomicName id="26876AB88F5DFFC7FF6CFA4C8B44DCD5" authority="(Johns et al. 1997)" baseAuthorityName="Johns" baseAuthorityYear="1997" box="[240,778,1422,1444]" class="Chondrichthyes" genus="Parvidiabolus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="longisulcus">
<emphasis id="D3F3CD298F5DFFC7FF6CFA4C8A53DCD2" box="[240,541,1422,1443]" italics="true" pageId="25" pageNumber="334">Parvidiabolus longisulcus</emphasis>
(
<bibRefCitation id="85166CCA8F5DFFC7FDA8FA4C8B4DDCD5" author="Johns MJ &amp; Barnes CR &amp; Orchard MJ" box="[564,771,1422,1444]" pageId="25" pageNumber="334" pagination="1 - 235" refId="ref19661" refString="Johns MJ, Barnes CR, Orchard MJ. 1997. Taxonomy and biostratigraphy of Middle and Late Triassic elasmobranch ichthyoliths from Northeastern British Columbia. Geological Survey of Canada, Bulletin 502: 1 - 235, plates 1 - 38." type="journal article" year="1997">
Johns 
<emphasis id="D3F3CD298F5DFFC7FD1AFA4D8AF8DCD2" box="[646,694,1422,1444]" italics="true" pageId="25" pageNumber="334">et al</emphasis>
. 1997
</bibRefCitation>
)
</taxonomicName>
from the Middle Triassic Liard Formation in western 
<collectingCountry id="999051AB8F5DFFC7FF3EFA0988B4DC90" box="[162,250,1483,1505]" name="Canada" pageId="25" pageNumber="334">Canada</collectingCountry>
(Ladinian). This material consists of scales only and does not settle the systematic position of these specimens. Although the histological ultrastructure observed in these scales is superficially similar to the ultrastructure of hybodont single-scale histology (see e.g. 
<bibRefCitation id="85166CCA8F5DFFC7FE91F9A789E1DF0B" author="Rieppel O" box="[269,431,1637,1659]" pageId="25" pageNumber="334" pagination="324 - 353" refId="ref20580" refString="Rieppel O. 1981. The hybodontiform sharks from the Middle Triassic of Mte. San Giorgio, Switzerland. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 161: 324 - 353." type="journal article" year="1981">Rieppel, 1981</bibRefCitation>
: fig 13E), we are hesitant to assign any systematic value to this feature. This 
<typeStatus id="3E3CAF998F5DFFC7FD44F9468B47DFE8" box="[728,777,1668,1689]" pageId="25" pageNumber="334">type</typeStatus>
of denticle is clearly distinctive from all other denticles found in patches of shark squamations from the Vega- Phroso Siltstone Member.
</paragraph>
<paragraph id="E138113B8F5DFFC7FF27F93C8CDBDF71" blockId="25.[162,779,441,1904]" lastBlockId="25.[826,1442,195,1536]" pageId="25" pageNumber="334">
The denticles described in TMP 97.74.10 (
<taxonomicName id="26876AB88F5DFFC7FF36F8DF890EDE43" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[170,320,1821,1842]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="aplopagus" status="gen. et sp. nov.">
<emphasis id="D3F3CD298F5DFFC7FF36F8DF890EDE43" box="[170,320,1821,1842]" italics="true" pageId="25" pageNumber="334">W. aplopagus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FED0F8DF89B1DE43" box="[332,511,1821,1842]" pageId="25" pageNumber="334" rank="species">gen. et sp. nov.</taxonomicNameLabel>
) exhibit extraordinary variation, even within a single dorsal fin, and there are several kinds of denticles on the entire body, but none is comparable to the denticle-type referred to as ‘ 
<emphasis id="D3F3CD298F5DFFC7FCA3FF208B87D986" box="[831,969,226,247]" italics="true" pageId="25" pageNumber="334">Palaeobates’</emphasis>
(compare 
<figureCitation id="79BC0DBE8F5DFFC7FBD8FF208CC7D986" box="[1092,1161,226,248]" captionStart="Figure 9" captionStartId="11.[162,240,1151,1170]" captionTargetBox="[162,1442,194,1119]" captionTargetId="figure-465@11.[162,1442,193,1119]" captionTargetPageId="11" captionText="Figure 9. Wapitiodus aplopagus gen. et sp. nov.: morphologic variation of dermal denticles as found in various body regions in holotype TMP 97.74.10. A, dermal denticles from between the pelvic and anal fins. B, dermal denticles from the tip of the lower jaw. C, dermal denticles from the dorsal trunk area. D, dermal denticles from the tip of the anterior dorsal fin. E, dermal denticles from the mid-area of the posterior dorsal fin. F, dermal denticles from the anterior rim of the fin above the posterior dorsal fin spine. The denticles are not necessarily either in situ or orientated in the same way, see the text for the description. The shark was drawn by Beat Scheffold, PIMUZ." figureDoi="http://doi.org/10.5281/zenodo.5428578" httpUri="https://zenodo.org/record/5428578/files/figure.png" pageId="25" pageNumber="334">Figs 9</figureCitation>
and 
<figureCitation id="79BC0DBE8F5DFFC7FB59FF208CBBD989" box="[1221,1269,226,248]" captionStart="Figure 19" captionStartId="21.[162,240,1165,1184]" captionTargetBox="[162,1442,194,1132]" captionTargetId="figure-406@21.[162,1442,193,1132]" captionTargetPageId="21" captionText="Figure 19. Ctenacanthoidea incertae sedis ‘genus A’: dermal denticles and unidentified elements as preserved in specimen UALVP 46572. A and B, bar-like?skeletal elements and impressions of elements aligned among numerous small denticles (arrows). C, denticle, and a sketch of the denticle, with diverging ridges in the crown. Possibly, this is the denticle-type referred to as ‘Palaeobates’ by Schaeffer &amp; Mangus (1976). D, thin section (UALVP 46572-T1) through one of the denticles, showing the osteodont pedicle and the mesial platform with orthodont prongs covered entirely with enameloid." figureDoi="http://doi.org/10.5281/zenodo.5428599" httpUri="https://zenodo.org/record/5428599/files/figure.png" pageId="25" pageNumber="334">19C</figureCitation>
). In particular, 
<bibRefCitation id="85166CCA8F5DFFC7FCA6FEC38C32D866" author="Schaeffer B &amp; Mangus M" box="[826,1148,257,279]" pageId="25" pageNumber="334" pagination="519 - 563" refId="ref20666" refString="Schaeffer B, Mangus M. 1976. An Early Triassic fish assemblage from British Columbia. Bulletin of the American Museum of Natural History 156: 519 - 563." type="journal article" year="1976">Schaeffer &amp; Mangus (1976)</bibRefCitation>
considered denticle morphology to be similar to 
<taxonomicName id="26876AB88F5DFFC7FBCBFEDD8B35D825" authority="(Stensio, 1921)" baseAuthorityName="Stensio" baseAuthorityYear="1921" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="polaris">
<emphasis id="D3F3CD298F5DFFC7FBCBFEDD8D7BD845" box="[1111,1333,287,308]" italics="true" pageId="25" pageNumber="334">Palaeobates polaris</emphasis>
(
<bibRefCitation id="85166CCA8F5DFFC7FADAFEDD8B3DD825" author="Stensio E" pageId="25" pageNumber="334" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö, 1921</bibRefCitation>
)
</taxonomicName>
. 
<bibRefCitation id="85166CCA8F5DFFC7FC10FEFC8C78D825" author="Stensio E" box="[908,1078,318,340]" pageId="25" pageNumber="334" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö (1921)</bibRefCitation>
, however, describes the denticles of 
<taxonomicName id="26876AB88F5DFFC7FC17FE9F8BBAD803" baseAuthorityName="Stensio" baseAuthorityYear="1921" box="[907,1012,349,370]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="polaris">
<emphasis id="D3F3CD298F5DFFC7FC17FE9F8BBAD803" box="[907,1012,349,370]" italics="true" pageId="25" pageNumber="334">P. polaris</emphasis>
</taxonomicName>
as ‘poorly preserved’ and states that a number of ridges extend backwards as ‘long slender processes’ (?ridges on mesial platform), but also states that the number of these ridges ‘... cannot be stated with certainty’. Nevertheless, as described in the 
<typeStatus id="3E3CAF998F5DFFC7FCF2FE348B81DB7D" box="[878,975,502,524]" pageId="25" pageNumber="334" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="26876AB88F5DFFC7FB9EFE358CD9DB7A" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[1026,1175,502,524]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="aplopagus" status="gen. et sp. nov.">
<emphasis id="D3F3CD298F5DFFC7FB9EFE358CD9DB7A" box="[1026,1175,502,524]" italics="true" pageId="25" pageNumber="334">W. aplopagus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FB3BFE358D26DB7D" box="[1191,1384,503,524]" pageId="25" pageNumber="334" rank="species">gen. et sp. nov.</taxonomicNameLabel>
, one given specimen may possess denticles of considerable variability, and even on the same fin it is possible to find denticles that have from none to numerous ridges in variably shaped platforms. The range of individual, intraspecific and interspecific variation in the denticles is rather unsatisfactorily known in Lower Triassic sharks, and it seems impossible at present to distinguish Early Mesozoic sharks on the basis of dermal denticle morphology alone (see also 
<bibRefCitation id="85166CCA8F5DFFC7FCA6FCEB8C0EDA4E" author="Mutter RJ &amp; Rieber H" box="[826,1088,809,831]" pageId="25" pageNumber="334" pagination="139 - 148" refId="ref20180" refString="Mutter RJ, Rieber H. 2005. Pyknotylacanthus spathianus gen. et sp. nov., a new ctenacanthoid from the Early Triassic of Bear Lake (Idaho, USA). Revista Brasileira de Paleontologia 8: 139 - 148." type="journal article" year="2005">Mutter &amp; Rieber, 2005</bibRefCitation>
). The assignment of any specimen from the Sulphur Mountain Formation to 
<taxonomicName id="26876AB88F5DFFC7FCA6FCA48BA6DA0D" box="[826,1000,870,892]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="D3F3CD298F5DFFC7FCA6FCA48BF1DA0A" box="[826,959,870,891]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
sp.
</taxonomicName>
is cast further into doubt in the light of the total evidence from this locality. There are several isolated teeth and a number of teeth preserved in the dentitions of the two most complete specimens of 
<taxonomicName id="26876AB88F5DFFC7FCC5FC238B97DA87" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[857,985,993,1014]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="D3F3CD298F5DFFC7FCC5FC238B97DA87" box="[857,985,993,1014]" italics="true" pageId="25" pageNumber="334">Wapitiodus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FC79FC238C03DA87" box="[997,1101,993,1014]" pageId="25" pageNumber="334" rank="genus">gen. nov.</taxonomicNameLabel>
, and none of these meet the criteria set out by 
<bibRefCitation id="85166CCA8F5DFFC7FB8BFC3D8CF3DD67" author="Stensio E" box="[1047,1213,1023,1046]" pageId="25" pageNumber="334" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö (1921)</bibRefCitation>
to describe 
<taxonomicName id="26876AB88F5DFFC7FAD3FC3D8B3ADD42" authorityName=": PIMUZ T" authorityYear="1179" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FAD3FC3D8B3ADD42" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
from Spitzbergen, i.e. ‘crown long and narrow, without lateral cones (cusps), but sometimes with principal cone’. A ‘longitudinal crista’ (i.e. ‘longitudinal ridge’ 
<emphasis id="D3F3CD298F5DFFC7FC33FBB98BA0DDE1" box="[943,1006,1147,1168]" italics="true" pageId="25" pageNumber="334">sensu</emphasis>
<bibRefCitation id="85166CCA8F5DFFC7FC6BFBB88C23DDFE" author="Reif W-E" box="[1015,1133,1146,1168]" pageId="25" pageNumber="334" pagination="39 - 55" refId="ref20554" refString="Reif W-E. 1973. Morphologie und Skulptur der Haifisch- Zahnkronen. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 143: 39 - 55." type="journal article" year="1973">Reif, 1973</bibRefCitation>
: fig. 2) is often present but may also be absent. These features are very vague and could (erroneously) be taken to be present in the teeth of 
<taxonomicName id="26876AB88F5DFFC7FC3AFB148C68DD9A" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[934,1062,1238,1259]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="D3F3CD298F5DFFC7FC3AFB148C68DD9A" box="[934,1062,1238,1259]" italics="true" pageId="25" pageNumber="334">Wapitiodus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FBA9FB158CD0DD9D" box="[1077,1182,1239,1260]" pageId="25" pageNumber="334" rank="genus">gen. nov.</taxonomicNameLabel>
In fact, the strongly asymmetric teeth with broad and flat crowns in 
<taxonomicName id="26876AB88F5DFFC7FCA6FAD18BF1DC59" authorityName=": PIMUZ T" authorityYear="1179" box="[826,959,1299,1320]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FCA6FAD18BF1DC59" box="[826,959,1299,1320]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
show an ornament consisting of faintly elevated and fine striae, sometimes either anastomosing or forming a network (
<bibRefCitation id="85166CCA8F5DFFC7FBEEFA938D5DDC17" author="Rieppel O" box="[1138,1299,1361,1383]" pageId="25" pageNumber="334" pagination="324 - 353" refId="ref20580" refString="Rieppel O. 1981. The hybodontiform sharks from the Middle Triassic of Mte. San Giorgio, Switzerland. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 161: 324 - 353." type="journal article" year="1981">Rieppel, 1981</bibRefCitation>
: fig. 9). Like the sectioned polyacrodontid tooth (TMP 88.98.51; 
<figureCitation id="79BC0DBE8F5DFFC7FCA6FA4C8BC6DCD5" box="[826,904,1422,1444]" captionStart="Figure 18" captionStartId="20.[142,220,989,1008]" captionTargetBox="[142,1422,194,956]" captionTargetId="figure-408@20.[142,1422,193,956]" captionTargetPageId="20" captionText="Figure 18. Morpho-histology of teeth (cf. Fig. 2). A, Wapitiodus aplopagus gen. et sp. nov.: an isolated tooth in labial view (holotype specimen TMP 97.74.10). B, Polyacrodontidae gen. et sp. indet.: vertical section through an isolated tooth, specimen TMP 88.98.51. Note that the root and most of the crown is osteodont. C–E, Polyacrodontidae gen. et sp. indet.: various scanning electron microscopy (SEM) magnifications of the SLE in specimen TMP 88.98.51 (white asterisks mark the same position in B–E). Abbreviations: ORT, orthodentine; OST, osteodentine; SLE, single layer enameloid." figureDoi="http://doi.org/10.5281/zenodo.5428597" httpUri="https://zenodo.org/record/5428597/files/figure.png" pageId="25" pageNumber="334">Fig. 18</figureCitation>
), the crowns of teeth of 
<taxonomicName id="26876AB88F5DFFC7FB2FFA4C8D76DCD2" authorityName=": PIMUZ T" authorityYear="1179" box="[1203,1336,1422,1443]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FB2FFA4C8D76DCD2" box="[1203,1336,1422,1443]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
are also covered by a layer of enameloid, but this layer is thicker in 
<taxonomicName id="26876AB88F5DFFC7FC2DFA098C7FDC91" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" box="[945,1073,1483,1504]" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="D3F3CD298F5DFFC7FC2DFA098C7FDC91" box="[945,1073,1483,1504]" italics="true" pageId="25" pageNumber="334">Wapitiodus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5DFFC7FBA6FA0E8CD1DC90" box="[1082,1183,1484,1505]" pageId="25" pageNumber="334" rank="genus">gen. nov.</taxonomicNameLabel>
, and almost the entire tooth consists of osteodentine.
</paragraph>
<paragraph id="E138113B8F5DFFC7FCA2F9FE8D7CDF02" blockId="25.[830,1435,1596,1652]" pageId="25" pageNumber="334">COMPARISON WITH LOWER TRIASSIC MATERIAL FROM THE?DIENERIAN OF SPITZBERGEN</paragraph>
<paragraph id="E138113B8F5DFFC4FCA6F9478A3AD8BE" blockId="25.[826,1442,1669,1905]" lastBlockId="26.[142,758,195,923]" lastPageId="26" lastPageNumber="335" pageId="25" pageNumber="334">
The lack of diagnostic skeletal elements associated with teeth in the sample from Wapiti Lake recalls the suspicion of 
<bibRefCitation id="85166CCA8F5DFFC7FC4FF9008CEADFA9" author="Stensio E" box="[979,1188,1730,1752]" pageId="25" pageNumber="334" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö (1921: 42)</bibRefCitation>
that the ‘generically indeterminable fin-spines’ from Spitzbergen, recovered as isolated fragments, may possibly be referred to 
<taxonomicName id="26876AB88F5DFFC7FCA6F8DC8B8DDE42" baseAuthorityName="Jaekel" baseAuthorityYear="1889" box="[826,963,1822,1843]" class="Chondrichthyes" family="Hybodontidae" genus="Polyacrodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FCA6F8DC8B8DDE42" box="[826,963,1822,1843]" italics="true" pageId="25" pageNumber="334">Polyacrodus</emphasis>
</taxonomicName>
(or 
<taxonomicName id="26876AB88F5DFFC7FC62F8DC8CCDDE42" authorityName=": PIMUZ T" authorityYear="1179" box="[1022,1155,1822,1843]" class="Chondrichthyes" family="Acrodontidae" genus="Palaeobates" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="25" pageNumber="334" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5DFFC7FC62F8DC8CCDDE42" box="[1022,1155,1822,1843]" italics="true" pageId="25" pageNumber="334">Palaeobates</emphasis>
</taxonomicName>
). The fin spines briefly described by 
<bibRefCitation id="85166CCA8F5DFFC7FC46F8FF8C9DDE22" author="Stensio E" box="[986,1235,1853,1875]" pageId="25" pageNumber="334" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö (1921: 40–42)</bibRefCitation>
resemble the fin spines from Wapiti Lake in various features, but most of these features can only be observed in very few and fragmentary specimens: stellate tubercles (all three of Stensiö’s specimens and all specimens with fin spines described here), broadly triangular shaped cross-section and convex posterior border (Stensiö’s specimen P.44 and UALVP 46528), the ‘enamel keel’ (Stensiö’s specimen P.35 and specimens TMP 97.74.10 and UALVP 46529) and the ultrastructure of the fin spines as far as is discernible (
<bibRefCitation id="85166CCA8F5EFFC4FE3EFE7B8A29D8BE" author="Stensio E" box="[418,615,441,463]" pageId="26" pageNumber="335" refId="ref20763" refString="Stensio E. 1921. Triassic Fishes from Spitzbergen. Wien, Adolf Holzhausen." type="book" year="1921">Stensiö, 1921: 41</bibRefCitation>
).
</paragraph>
<paragraph id="E138113B8F5EFFC4FF3BFE1A893EDB38" blockId="26.[142,758,195,923]" pageId="26" pageNumber="335">
In 1932, Stensiö described a few more teeth on which he erected 
<taxonomicName id="26876AB88F5EFFC4FECAFE358A5CDB7A" baseAuthorityName="Stensio" baseAuthorityYear="1932" box="[342,530,502,524]" class="Chondrichthyes" family="Hybodontidae" genus="Polyacrodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="26" pageNumber="335" phylum="Chordata" rank="species" species="claveringensis">
<emphasis id="D3F3CD298F5EFFC4FECAFE358A5CDB7A" box="[342,530,502,524]" italics="true" pageId="26" pageNumber="335">P. claveringensis</emphasis>
</taxonomicName>
and four additional fragmentary fin spines, two of which he called ‘ 
<taxonomicName id="26876AB88F5EFFC4FF0FFDF1897EDB39" box="[147,304,563,584]" class="Chondrichthyes" family="Hybodontidae" genus="Nemacanthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hybodontiformes" pageId="26" pageNumber="335" phylum="Chordata" rank="genus">
<emphasis id="D3F3CD298F5EFFC4FF0FFDF1897EDB39" box="[147,304,563,584]" italics="true" pageId="26" pageNumber="335">Nemacanthus</emphasis>
</taxonomicName>
-like’.
</paragraph>
<paragraph id="E138113B8F5EFFC4FF3BFD908A15DAEA" blockId="26.[142,758,195,923]" pageId="26" pageNumber="335">
<bibRefCitation id="85166CCA8F5EFFC4FF3BFD908902DB19" author="Stensio E" box="[167,332,594,616]" pageId="26" pageNumber="335" pagination="1 - 298" refId="ref20778" refString="Stensio E. 1932. Triassic fishes from East Greenland. Meddelelser Om Gronland 83: 1 - 298, plates 1 - 39." type="journal article" year="1932">Stensiö (1932)</bibRefCitation>
also reported dermal denticles from the head region of 
<taxonomicName id="26876AB88F5EFFC4FEE3FDB38AF9DBF7" baseAuthorityName="Stensio" baseAuthorityYear="1932" box="[383,695,625,646]" class="Chondrichthyes" family="Hybodontidae" genus="Polyacrodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="26" pageNumber="335" phylum="Chordata" rank="species" species="claveringensis">
<emphasis id="D3F3CD298F5EFFC4FEE3FDB38AF9DBF7" box="[383,695,625,646]" italics="true" pageId="26" pageNumber="335">Polyacrodus claveringensis</emphasis>
</taxonomicName>
that actually resemble our 
<figureCitation id="79BC0DBE8F5EFFC4FE17FD4D89B8DBD4" box="[395,502,655,677]" captionStart="Figure 9" captionStartId="11.[162,240,1151,1170]" captionTargetBox="[162,1442,194,1119]" captionTargetId="figure-465@11.[162,1442,193,1119]" captionTargetPageId="11" captionText="Figure 9. Wapitiodus aplopagus gen. et sp. nov.: morphologic variation of dermal denticles as found in various body regions in holotype TMP 97.74.10. A, dermal denticles from between the pelvic and anal fins. B, dermal denticles from the tip of the lower jaw. C, dermal denticles from the dorsal trunk area. D, dermal denticles from the tip of the anterior dorsal fin. E, dermal denticles from the mid-area of the posterior dorsal fin. F, dermal denticles from the anterior rim of the fin above the posterior dorsal fin spine. The denticles are not necessarily either in situ or orientated in the same way, see the text for the description. The shark was drawn by Beat Scheffold, PIMUZ." figureDoi="http://doi.org/10.5281/zenodo.5428578" httpUri="https://zenodo.org/record/5428578/files/figure.png" pageId="26" pageNumber="335">Figures 9</figureCitation>
(B–D) and 12. As mentioned above, there are no anteriormost teeth preserved and the lateral/postero-lateral teeth retain a small central cusp. Two of the 
<bibRefCitation id="85166CCA8F5EFFC4FDBFFD298A82DA70" author="Stensio E" box="[547,716,747,769]" pageId="26" pageNumber="335" pagination="1 - 298" refId="ref20778" refString="Stensio E. 1932. Triassic fishes from East Greenland. Meddelelser Om Gronland 83: 1 - 298, plates 1 - 39." type="journal article" year="1932">Stensiö (1932)</bibRefCitation>
fin spines (nrs 2 and 3) show a tubercular ornament and an enamel keel (cf. 
<figureCitation id="79BC0DBE8F5EFFC4FEEDFCEB898CDA4E" box="[369,450,809,831]" captionStart="Figure 8" captionStartId="10.[142,220,1250,1269]" captionTargetBox="[142,1438,194,1218]" captionTargetId="figure-423@10.[142,1438,194,1218]" captionTargetPageId="10" captionText="Figure 8. Wapitiodus homalorhizo sp. nov.: structure of fin spine in cross-sections (A–D) through the apical half of specimen UALVP 46528 (thin sections T1–3) and the external view of the apex of the fin spine in specimen UALVP 46529 (E, silicon peel dusted with NH4Cl). A, overview: note the large but secondarily obliterated, partly remodeled cavity and the large tubercles clearly delimited from the core of the fin spine by lines of arrested growth. B, note the couple of displaced and stacked tubercles in the left postero-lateral corner (as indicated in A) at a slightly lower level of the spine (thin section T3). C, note the internal structure in the posterolateral wall of the fin spine that is characteristic of ctenacanthoids. D, attempts at restoration of the cross-section through the apical third. E, note the well-spaced stellate tubercles (arrowhead 2), the absence of recurved denticles in the posterolateral wall (arrowhead 1, but also see Fig. 10 and the description of specimen UALVP 46528) and the partly preserved anterior rim (arrowhead 3)." figureDoi="http://doi.org/10.5281/zenodo.5428576" httpUri="https://zenodo.org/record/5428576/files/figure.png" pageId="26" pageNumber="335">Fig. 8E</figureCitation>
) that are similar to 
<taxonomicName id="26876AB88F5EFFC4FD2FFCEB8901DA2D" authority="Mutter &amp; Blanger &amp; Neuman, 2007" authorityName="Mutter &amp; Blanger &amp; Neuman" authorityYear="2007" class="Chondrichthyes" family="Homalodontidae" genus="Wapitiodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Selachii" pageId="26" pageNumber="335" phylum="Chordata" rank="species" species="aplopagus" status="gen. et sp. nov.">
<emphasis id="D3F3CD298F5EFFC4FD2FFCEB8901DA2D" italics="true" pageId="26" pageNumber="335">Wapitiodus aplopagus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="C8C070528F5EFFC4FECAFC8A89B4DA2C" box="[342,506,840,861]" pageId="26" pageNumber="335" rank="species">gen. et sp. nov.</taxonomicNameLabel>
Because of the imperfect state of preservation of these remains, however, these finds cannot be further compared.
</paragraph>
</subSubSection>
</treatment>
</document>