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<emphasis id="2EA8EAF6667DFFC2FE8317B4F8CFFC4B" bold="true" box="[330,454,940,963]" italics="true" pageId="8" pageNumber="86">Theatonius</emphasis>
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<paragraph id="1C6336E4667DFFC2FE9117C4F8F3FC7A" blockId="8.[344,506,988,1010]" box="[344,506,988,1010]" pageId="8" pageNumber="86">
(
<figureCitation id="84E72A61667DFFC2FE9617C4F8FAFC7A" box="[351,499,988,1010]" captionStart="Text-fig" captionStartId="8.[106,187,616,637]" captionTargetBox="[106,1459,152,574]" captionTargetId="figure-607@8.[121,1447,159,558]" captionTargetPageId="8" captionText="Text-fig. 5. Edentulous anuran maxillae from the middle late Campanian (Judithian) of Alberta, Canada and Utah, USA. All images are photographs and depict specimens lightly dusted with ammonium chloride to enhance details and texture. Images at different magnifications; see corresponding scale bars. af Tyrrellbatrachus brinkmani GARDNER, 2015, all from basal part of Dinosaur Park Formation, Dinosaur Provincial Park,Alberta: a, b incomplete right maxilla, TMP 1985.066.0035 (holotype), in labial (a) and lingual (b) views, from TMP locality L0404; c, d incomplete right maxilla, TMP 1986.033.0033, in labial (c) and lingual (d) views, from TMP locality L0031; e, f incomplete left maxilla, TMP 1986.214.0032, in labial (e) and lingual (f) views, from TMP locality L0051. g, h Theatonius n. sp., nearly complete left maxilla, OMNH 67082, in labial (g) and lingual (h) views, from Kaiparowits Formation, OMNH locality V6, Utah." figureDoi="http://doi.org/10.5281/zenodo.4773074" httpUri="https://zenodo.org/record/4773074/files/figure.png" pageId="8" pageNumber="86">
Text-fig.
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, h
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)
</paragraph>
<paragraph id="1C6336E4667DFFC2FF591011FBE0FBA8" blockId="8.[106,745,1033,1087]" box="[144,745,1033,1056]" pageId="8" pageNumber="86">M a t e r i a l a n d o c c u r r e n c e s: Six maxillae from</paragraph>
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Kaiparowits Formation,
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,
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(Appendix 2).
</paragraph>
<paragraph id="1C6336E4667DFFC2FF591042F8EEF828" blockId="8.[106,745,1114,1952]" pageId="8" pageNumber="86">
D e s c r i p t i o n: The six maxillary specimens inclu- de a fragmentary example from the UMNH collections previously figured as being from an unidentified anuran (
<bibRefCitation id="784D4B15667DFFC2FFB810A0F84EFB47" author="Rocek, Z. &amp; Eaton, J. G. &amp; Gardner, J. D. &amp; Prikryl, T." box="[113,327,1208,1231]" pageId="8" pageNumber="86" pagination="341 - 394" refId="ref26694" refString="Rocek, Z., Eaton, J. G., Gardner, J. D., Prikryl, T. (2010): Evolution of anuran assemblages in the Late Cretaceous of Utah, USA. - Palaeobiodiversity and Palaeoenvironments, 90 (4): 341 - 394. http: // dx. doi. org / 10.1007 / s 12549 - 010 - 0040 - 2" type="journal article" year="2010">Roček et al. 2010</bibRefCitation>
: fig. 16Ab) plus five more recently identified examples from the OMNH collections: a nearly complete left maxilla (OMNH 67082;
<figureCitation id="84E72A61667DFFC2FDDB10EFFBBBFA86" box="[530,690,1271,1294]" captionStart="Text-fig" captionStartId="8.[106,187,616,637]" captionTargetBox="[106,1459,152,574]" captionTargetId="figure-607@8.[121,1447,159,558]" captionTargetPageId="8" captionText="Text-fig. 5. Edentulous anuran maxillae from the middle late Campanian (Judithian) of Alberta, Canada and Utah, USA. All images are photographs and depict specimens lightly dusted with ammonium chloride to enhance details and texture. Images at different magnifications; see corresponding scale bars. af Tyrrellbatrachus brinkmani GARDNER, 2015, all from basal part of Dinosaur Park Formation, Dinosaur Provincial Park,Alberta: a, b incomplete right maxilla, TMP 1985.066.0035 (holotype), in labial (a) and lingual (b) views, from TMP locality L0404; c, d incomplete right maxilla, TMP 1986.033.0033, in labial (c) and lingual (d) views, from TMP locality L0031; e, f incomplete left maxilla, TMP 1986.214.0032, in labial (e) and lingual (f) views, from TMP locality L0051. g, h Theatonius n. sp., nearly complete left maxilla, OMNH 67082, in labial (g) and lingual (h) views, from Kaiparowits Formation, OMNH locality V6, Utah." figureDoi="http://doi.org/10.5281/zenodo.4773074" httpUri="https://zenodo.org/record/4773074/files/figure.png" pageId="8" pageNumber="86">
Text-fig.
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, h
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) and four less complete maxillae (unfigured). Collectively these
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document the entire structure of the maxilla. These specimens are strikingly similar to maxillae of the late Maastrichtian (Lancian) anuran
<taxonomicName id="DBDC4D67667DFFC2FE04116CFBBDFA03" authorityName="FOX" authorityYear="1976" box="[461,692,1396,1419]" class="Amphibia" family="Tregobatrachidae" genus="Theatonius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="8" pageNumber="86" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="2EA8EAF6667DFFC2FE04116CFBBDFA03" box="[461,692,1396,1419]" italics="true" pageId="8" pageNumber="86">Theatonius lancensis</emphasis>
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(see revised species diagnosis by
<bibRefCitation id="784D4B15667DFFC2FE54118CFB38FA23" author="Gardner, J. D." box="[413,561,1428,1451]" pageId="8" pageNumber="86" pagination="219 - 249" refId="ref24635" refString="Gardner, J. D. (2008): New information on frogs (Lissamphibia: Anura) from the Lance Formation (late Maastrichtian) and Bug Creek Anthills (late Maastrichtian and early Paleoecene), Hell Creek Formation, USA. - In: Sankey, J. T., Baszio, B. (eds), Vertebrate Microfossil Assemblages: Their Role in Paleoecology and Paleobiogeography, Indiana University Press, Bloomington, pp. 219 - 249." type="book chapter" year="2008">Gardner 2008</bibRefCitation>
) in the following features: small size (OMNH 67082 is 4.9 mm long); labial surface ornamented with moderate-sized and closely packed pustules; crista dentalis moderately deep and lacks teeth; lamina horizontalis a weakly developed and lingually convex ridge; margo orbitalis deeply and asymmetrically concave; preorbital area taller and longer than postorbital area; processus palatinus relatively massive, projects linguodorsally, and dorsally bears a prominent facet; groove for ductus nasolacrimalis deep, wraps around labial base of processus palatinus and extends onto margo orbitalis; anterior portion of lamina anterior lingually bears a prominent facet; rostellum small, pointed, and directed anteriorly; processus pterygoideus lingually short, but anteroposteriorly elongate, and bears prominent articular facet that wraps posterodorsally onto processus zygomatico-maxillaris; and posterior end of processus posterior bluntly pointed.
</paragraph>
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R e m a r k s: As indicated above, maxillae reported here from the Kaiparowits Formation compare favourably with the type species
<taxonomicName id="DBDC4D67667DFFC2FEE613E4FB13F79B" authorityName="FOX" authorityYear="1976" box="[303,538,2044,2067]" class="Amphibia" family="Tregobatrachidae" genus="Theatonius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="8" pageNumber="86" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="2EA8EAF6667DFFC2FEE613E4FB13F79B" box="[303,538,2044,2067]" italics="true" pageId="8" pageNumber="86">Theatonius lancensis</emphasis>
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. The one notable difference is that the occlusolingual rim of the pars dentalis bears a row of tiny bumps in the
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specimens, whereas that margin is smooth in the two maxillae (
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and one referred:
<bibRefCitation id="784D4B15667DFFC2FC5017F4FD3DFB8B" author="Gardner, J. D." box="[921,1076,1004,1027]" pageId="8" pageNumber="86" pagination="219 - 249" refId="ref24635" refString="Gardner, J. D. (2008): New information on frogs (Lissamphibia: Anura) from the Lance Formation (late Maastrichtian) and Bug Creek Anthills (late Maastrichtian and early Paleoecene), Hell Creek Formation, USA. - In: Sankey, J. T., Baszio, B. (eds), Vertebrate Microfossil Assemblages: Their Role in Paleoecology and Paleobiogeography, Indiana University Press, Bloomington, pp. 219 - 249." type="book chapter" year="2008">Gardner 2008</bibRefCitation>
: fig. 13.2AD and E, respectively) described for
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<emphasis id="2EA8EAF6667DFFC2FC2F1014FDD6FBAB" box="[998,1247,1036,1059]" italics="true" pageId="8" pageNumber="86">Theatonius lancensis</emphasis>
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from the Lance Formation. That morphological difference, coupled with differences in ages (i.e., middle late Campanian vs. late Maastrichtian), suggest the maxillae from the Kaiparowits Formation pertain to a second species of
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<emphasis id="2EA8EAF6667DFFC2FAFA1094FCA5FB2B" box="[1331,1452,1164,1187]" italics="true" pageId="8" pageNumber="86">Theatonius</emphasis>
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. Unfortunately, no examples of the equally distinctive frontoparietals or squamosals comparable to those known for
<taxonomicName id="DBDC4D67667DFFC2FC9010F4FADCFA8B" authorityName="FOX" authorityYear="1976" box="[857,981,1260,1283]" class="Amphibia" family="Tregobatrachidae" genus="Theatonius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="8" pageNumber="86" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="2EA8EAF6667DFFC2FC9010F4FADCFA8B" box="[857,981,1260,1283]" italics="true" pageId="8" pageNumber="86">T. lancensis</emphasis>
</taxonomicName>
(see
<bibRefCitation id="784D4B15667DFFC2FBC510F4FD88FA8B" author="Fox, R. C." box="[1036,1153,1260,1283]" pageId="8" pageNumber="86" pagination="1486 - 1490" refId="ref24212" refString="Fox, R. C. (1976 b): An edentulous frog (Theatonius lancensis, new genus and species) from the Upper Cretaceous Lance Formation of Wyoming. - Canadian Journal of Earth Sciences, 13 (10): 1486 - 1490. http: // dx. doi. org / 10.1139 / e 76 - 154" type="journal article" year="1976">Fox 1976b</bibRefCitation>
,
<bibRefCitation id="784D4B15667DFFC2FB4210F4FC28FA8B" author="Gardner, J. D." box="[1163,1313,1260,1283]" pageId="8" pageNumber="86" pagination="219 - 249" refId="ref24635" refString="Gardner, J. D. (2008): New information on frogs (Lissamphibia: Anura) from the Lance Formation (late Maastrichtian) and Bug Creek Anthills (late Maastrichtian and early Paleoecene), Hell Creek Formation, USA. - In: Sankey, J. T., Baszio, B. (eds), Vertebrate Microfossil Assemblages: Their Role in Paleoecology and Paleobiogeography, Indiana University Press, Bloomington, pp. 219 - 249." type="book chapter" year="2008">Gardner 2008</bibRefCitation>
: fig. 13.2FJ) have been identified among the
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samples.
</paragraph>
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The occurrence of
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<emphasis id="2EA8EAF6667DFFC2FB811133FDC8FACA" box="[1096,1217,1323,1346]" italics="true" pageId="8" pageNumber="86">Theatonius</emphasis>
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in the Kaiparowits Formation of south-central
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extends the geographical range for the genus southwards into
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from
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(
<bibRefCitation id="784D4B15667DFFC2FCF51191FAB2FA28" author="Fox, R. C." box="[828,955,1417,1440]" pageId="8" pageNumber="86" pagination="1486 - 1490" refId="ref24212" refString="Fox, R. C. (1976 b): An edentulous frog (Theatonius lancensis, new genus and species) from the Upper Cretaceous Lance Formation of Wyoming. - Canadian Journal of Earth Sciences, 13 (10): 1486 - 1490. http: // dx. doi. org / 10.1139 / e 76 - 154" type="journal article" year="1976">Fox 1976b</bibRefCitation>
) and
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(
<bibRefCitation id="784D4B15667DFFC2FBB51191FCA6FA28" author="Gardner, J. D. &amp; DeMar, D. G., Jr." box="[1148,1455,1417,1440]" pageId="8" pageNumber="86" pagination="459 - 515" refId="ref24890" refString="Gardner, J. D., DeMar, D. G., Jr. (2013): Mesozoic and Paleocene lissamphibian assemblages of North America: a comprehensive review. - Palaeobiodiversity and Palaeoenvironments, 93 (4): 459 - 515. http: // dx. doi. org / 10.1007 / s 12549 - 013 - 0130 - z" type="journal article" year="2013">Gardner and DeMar 2013</bibRefCitation>
,
<bibRefCitation id="784D4B15667DFFC2FCFD11B0FAF3FA37" author="Mercier, G. &amp; DeMar, D. G., Jr. &amp; Wilson, G. P." box="[820,1018,1448,1471]" pageId="8" pageNumber="86" pagination="187" refId="ref25915" refString="Mercier, G., DeMar, D. G., Jr., Wilson, G. P. (2014): Frogs and toads (Lissamphibia, Anura) during the end-Cretaceous mass extinction: evidence from the fossil record of northeastern Montana. - In: Program and Abstracts, 74 th Annual Meeting, Society of Vertebrate Paleontology, p. 187." type="book chapter" year="2014">Mercier et al. 2014</bibRefCitation>
) and extends its temporal range back from the late Maastrichtian to the middle late Campanian. There also is an unconfirmed report of
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<emphasis id="2EA8EAF6667DFFC2FB4211FEFC0BFA75" box="[1163,1282,1510,1533]" italics="true" pageId="8" pageNumber="86">Theatonius</emphasis>
</taxonomicName>
-like squamosals in the Campanian (possibly Aquilan or Judithian equivalent; see
<bibRefCitation id="784D4B15667DFFC2FCAD123DFD3EF9B4" author="Cifelli, R. L. &amp; Eberle, J. J. &amp; Lofgren, D. L. &amp; Lillegraven, J. A. &amp; Clemens, W. A." box="[868,1079,1573,1596]" pageId="8" pageNumber="86" pagination="21 - 42" refId="ref22191" refString="Cifelli, R. L., Eberle, J. J., Lofgren, D. L., Lillegraven, J. A., Clemens, W. A. (2004): Mammalian biochronology of the latest Cretaceous. - In: Woodburne, M. O. (ed.), Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology, Columbia University Press, New York, pp. 21 - 42." type="book chapter" year="2004">Cifelli et al. 2004</bibRefCitation>
,
<bibRefCitation id="784D4B15667DFFC2FB8E123DFCA5F9B4" author="Kielan-Jaworowska, Z. &amp; Cifelli, R. L. &amp; Luo, Z. - X." box="[1095,1452,1573,1596]" pageId="8" pageNumber="86" refId="ref25705" refString="Kielan-Jaworowska, Z., Cifelli, R. L., Luo, Z. - X. (2004): Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure. - Columbia University Press, New York, 630 pp." type="book" year="2004">Kielan-Jaworowska et al. 2004</bibRefCitation>
) Marshalltown Formation of
<collectingRegion id="DE18F806667DFFC2FB92125CFDDCF9D3" box="[1115,1237,1604,1627]" country="United States of America" name="New Jersey" pageId="8" pageNumber="86">New Jersey</collectingRegion>
(
<bibRefCitation id="784D4B15667DFFC2FB28125CFA67F9F2" author="Denton, R. K., Jr. &amp; O'Neill, R. C." pageId="8" pageNumber="86" pagination="484 - 494" refId="ref22816" refString="Denton, R. K., Jr., O'Neill, R. C. (1998): Parrisia neocesariensis, a new batrachosauroidid salamander and other amphibians from the Campanian of eastern North America. - Journal of Vertebrate Paleontology, 18 (3): 484 - 494. http: // dx. doi. org / 10.1080 / 02724634.1998.10011076" type="journal article" year="1998">Denton and ONeill 1998</bibRefCitation>
). Compared to the apparently restricted distribution of
<taxonomicName id="DBDC4D67667DFFC2FC9A129AFADDF911" authorityName="FOX" authorityYear="1976" box="[851,980,1666,1689]" class="Amphibia" family="Tregobatrachidae" genus="Theatonius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="8" pageNumber="86" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="2EA8EAF6667DFFC2FC9A129AFADDF911" box="[851,980,1666,1689]" italics="true" pageId="8" pageNumber="86">T. lancensis</emphasis>
</taxonomicName>
in the Lance Formation, where it is known only from the Bushy Tailed Blowout locality, the putative new species of
<taxonomicName id="DBDC4D67667DFFC2FC3012D9FD7BF950" authorityName="Gardner &amp; Redman &amp; Cifelli" authorityYear="2016" box="[1017,1138,1729,1752]" class="Amphibia" family="Tregobatrachidae" genus="Theatonius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="8" pageNumber="86" phylum="Chordata" rank="genus">
<emphasis id="2EA8EAF6667DFFC2FC3012D9FD7BF950" box="[1017,1138,1729,1752]" italics="true" pageId="8" pageNumber="86">Theatonius</emphasis>
</taxonomicName>
from
<collectingRegion id="DE18F806667DFFC2FB0112D9FDF4F950" box="[1224,1277,1729,1752]" country="United States of America" name="Utah" pageId="8" pageNumber="86">Utah</collectingRegion>
has a broader stratigraphic distribution, being represented at five localities in both the lower (OMNH V6 and V9; UMNH VP 108) and upper (OMNH V5 and 61) parts of the Kaiparowits Formation.
</paragraph>
</subSubSection>
</treatment>
</document>