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<document id="5E3ABDE46229B4AE53E7591E6DC4E94A" ID-ISSN="0024-4082" ID-ZooBank="E79B2B6DBB6A-4DD2-A291-A2563541885A" IM.materialsCitations_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630107096780" checkinUser="felipe" docAuthor="Cossette, Adam P." docDate="2021" docId="03839C7AC4207F25BD9CAD55FADBFA6E" docLanguage="en" docName="zlz178.pdf" docOrigin="Zoological Journal of the Linnean Society 191" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Bottosaurus fustidens Cossette 2021, SP. NOV." docType="treatment" docVersion="2" masterDocId="FFBAE402C4237F21BD6FAB63FFC9FF87" masterDocTitle="A new species of Bottosaurus (Alligatoroidea: Caimaninae) from the Black Peaks Formation (Palaeocene) of Texas indicates an early radiation of North American caimanines" updateTime="1699154177583" updateUser="plazi">
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<mods:title id="2CC4A2354EF145EF19E879F0635A885E">A new species of Bottosaurus (Alligatoroidea: Caimaninae) from the Black Peaks Formation (Palaeocene) of Texas indicates an early radiation of North American caimanines</mods:title>
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<mods:namePart id="E9288AF220D55640A27489C8CF844F01">Cossette, Adam P.</mods:namePart>
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<mods:title id="10D6405DB1D764C2728406A3FBD08995">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="6B9B66C1F43BD2302EA6BB600D332E63">2021</mods:date>
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<treatment id="03839C7AC4207F25BD9CAD55FADBFA6E" LSID="urn:lsid:plazi:treatment:03839C7AC4207F25BD9CAD55FADBFA6E" httpUri="http://treatment.plazi.org/id/03839C7AC4207F25BD9CAD55FADBFA6E" lastPageId="4" pageId="3">
<subSubSection id="C3307EE7C4207F22BD9CAD55FE36F915" pageId="3" type="nomenclature">
<paragraph id="8B952D6CC4207F22BD9CAD55FD5FF9CA" blockId="3.[243,662,1589,1613]" box="[243,662,1589,1613]" pageId="3">
<heading id="D0DD9A00C4207F22BD9CAD55FD5FF9CA" box="[243,662,1589,1613]" centered="true" fontSize="9" level="2" pageId="3" reason="2">
<taxonomicName id="4C2A56EFC4207F22BD9CAD55FDE4F9CB" authority="Cossette, 2021" authorityName="Cossette" authorityYear="2021" box="[243,557,1590,1613]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="3" phylum="Chordata" rank="species" species="fustidens" status="sp. nov.">BOTTOSAURUS FUSTIDENS</taxonomicName>
<emphasis id="B95EF17EC4207F22BF5BAD5AFD5FF9CA" bold="true" box="[564,662,1589,1613]" pageId="3">
<taxonomicNameLabel id="A26D4C05C4207F22BF5BAD5AFD5FF9CA" box="[564,662,1589,1613]" pageId="3" rank="species">SP. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B952D6CC4207F22BDFEAD3DFE36F915" blockId="3.[145,761,1630,1682]" pageId="3">
l s i d: u r n: l s i d: z o o b a n k. o r g: a c t:
<uuid id="FF8C17B9C4207F22BF38AD3DFE36F915" pageId="3">9 F 6 E B 6 0 1 - 1913-4421-9562-1652A376A89A</uuid>
</paragraph>
</subSubSection>
<caption id="DF557DE4C4207F22BE47AD3DFB3BF8A3" pageId="3" startId="3.[808,888,1630,1652]" targetBox="[808,1425,1283,1589]" targetPageId="3">
<paragraph id="8B952D6CC4207F22BE47AD3DFB3BF8A3" blockId="3.[806,1422,1630,1828]" pageId="3">
<emphasis id="B95EF17EC4207F22BE47AD3DFC5BF9F4" bold="true" box="[808,914,1630,1652]" pageId="3">Figure 2.</emphasis>
TMM 40148-7, holotype,
<taxonomicName id="4C2A56EFC4207F22B9C9AD3CFA47F9F3" authorityName="Cossette" authorityYear="2021" box="[1190,1422,1630,1652]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="3" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4207F22B9C9AD3CFA47F9F3" box="[1190,1422,1630,1652]" italics="true" pageId="3">Bottosaurus fustidens</emphasis>
</taxonomicName>
and NJSM 11265,
<taxonomicName id="4C2A56EFC4207F22BE94AD1FFB16F916" authorityName="Meyer" authorityYear="1832" box="[1019,1247,1659,1681]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="3" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4207F22BE94AD1FFB16F916" box="[1019,1247,1659,1681]" italics="true" pageId="3">Bottosaurus harlani</emphasis>
</taxonomicName>
, comparison of jugal region in dorsal view. A, TMM 40148-7, holotype,
<taxonomicName id="4C2A56EFC4207F22BE49ADD4FC6AF94B" authorityName="Cossette" authorityYear="2021" box="[806,931,1718,1740]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="3" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4207F22BE49ADD4FC6AF94B" box="[806,931,1718,1740]" italics="true" pageId="3">B. fustidens</emphasis>
</taxonomicName>
right ascending process of jugal and anterior margin of medial jugal foramen. B, NJSM 11265,
<taxonomicName id="4C2A56EFC4207F22B84BADB7FA47F96E" authorityName="Meyer" authorityYear="1832" box="[1316,1422,1747,1769]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="3" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4207F22B84BADB7FA47F96E" box="[1316,1422,1747,1769]" italics="true" pageId="3">B. harlani</emphasis>
</taxonomicName>
left jugal. Abbreviations: apj, ascending process of jugal; mjf, medial jugal foramen. Scale bar = 5 cm.
</paragraph>
</caption>
<subSubSection id="C3307EE7C4207F25BDFEADD2FADBFA6E" lastPageId="4" pageId="3" type="description">
<paragraph id="8B952D6CC4207F25BDFEADD2FDAFFA2C" blockId="3.[145,761,1712,1826]" lastBlockId="4.[163,779,908,1451]" lastPageId="4" pageId="3">
<emphasis id="B95EF17EC4207F22BDFEADD2FEDAF941" box="[145,275,1712,1734]" italics="true" pageId="3">H o l o t y p e:</emphasis>
T M M 4 0 1 4 8 - 7, i n c o m p l e t e s k u l l, including the right posterior dorsal process of the premaxilla, mostly complete nasals, frontal, prefrontals, mostly complete postorbitals, anterior parietal, mostly complete squamosals, partial jugals, incomplete quadratojugals, an incomplete left quadrate, portions of the right and left exoccipitals, partial ectopterygoids and dorsal portions of the pterygoids. Nearly complete lower jaws are preserved. The right dentary is more complete than the left. Both splenials are preserved, and the right side is nearly complete. The anterior left angular and the posterior right angular are preserved; the right posterior surangular is present, and most of the left articular is preserved. Postcranial elements are preserved. Four cervical vertebrae, six dorsal vertebrae, six caudal vertebrae and four indeterminate vertebral fragments are preserved; all vertebrae are incomplete. Portions of dorsal ribs are poorly preserved. The left humerus is incomplete. The proximal right ischium is preserved. The left femur is complete. Numerous elements of the hand and foot are preserved (
<figureCitation id="131131E9C4277F25BC9AAEF6FD9EFA2C" box="[501,599,1429,1451]" captionStart-0="Figure 4" captionStart-1="Figure 5" captionStart-2="Figure 6" captionStartId-0="6.[163,243,1795,1817]" captionStartId-1="8.[162,240,1522,1544]" captionStartId-2="10.[163,241,1734,1756]" captionTargetBox-0="[219,1386,192,1756]" captionTargetBox-1="[322,1282,197,1483]" captionTargetBox-2="[286,1320,194,1694]" captionTargetId-0="figure-24@6.[219,1386,192,1756]" captionTargetId-1="figure-23@8.[322,1282,197,1483]" captionTargetId-2="figure-25@10.[286,1320,194,1694]" captionTargetPageId-0="6" captionTargetPageId-1="8" captionTargetPageId-2="10" captionText-0="Figure 4. TMM 40148-7, holotype, Bottosaurus fustidens posterior skull and snout. A, snout and anterior skull table in dorsal view. B, posterior left maxilla and anterior jugal in dorsal view. C, left pterygoid and ectopterygoid in anterior and posterior views. D, left posterior jugal and anterior quadratojugal. E, left posterior quadratojugal and quadrate in dorsal view. F, left squamosal and quadrate in lateral view. G, left postorbital and squamosal in dorsal view. H, right postorbital" captionText-1="Figure 5. TMM 40148-7, holotype, Bottosaurus fustidens lower jaw. AC, right hemimandible in dorsal (A), medial (B) and lateral (C) views. DG, left posterior mandible in dorsal (D), lateral (E), medial (F) and ventral (G) views. Abbreviations: alv, alveoli; an, angular; ar, articular; d, dentary; d#, numbered dentary tooth; fio, foramen intermandibularis oralis; ms, mandibular symphysis; s, splenial; sa, surangular. Scale bar = 5 cm." captionText-2="Figure 6. TMM 40148-7, holotype, Bottosaurus fustidens selected postcrania. A, left femur in lateral view. B, left humerus in lateral view. C, right proximal ischium in medial view. D, metatarsal I in ventral view. E, ungule phalanx in lateral view. F, axis in lateral, dorsal and anterior views. G, thoracic vertebra in posterior and lateral views. H, thoracic vertebra in posterior and lateral views. I, thoracic vertebra in posterior and lateral views. J, thoracic vertebra in posterior and lateral views. K, caudal vertebra in posterior and lateral views. L, caudal vertebra in posterior and lateral views. M, caudal vertebra in posterior and lateral views. Abbreviation: op, odontoid process. Scale bar = 5 cm." pageId="4">Figs 46</figureCitation>
).
</paragraph>
<caption id="DF557DE4C4277F25BDCCA86CFE9FFCE7" pageId="4" startId="4.[163,242,783,805]" targetBox="[323,1283,195,744]" targetPageId="4">
<paragraph id="8B952D6CC4277F25BDCCA86CFE9FFCE7" blockId="4.[163,1443,783,864]" pageId="4">
<emphasis id="B95EF17EC4277F25BDCCA86CFEC5FCA3" bold="true" box="[163,268,783,805]" pageId="4">Figure 3.</emphasis>
Comparison of
<taxonomicName id="4C2A56EFC4277F25BCD8A873FD54FCA2" authorityName="Cossette" authorityYear="2021" box="[439,669,783,805]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4277F25BCD8A873FD54FCA2" box="[439,669,783,805]" italics="true" pageId="4">Bottosaurus fustidens</emphasis>
</taxonomicName>
and
<taxonomicName id="4C2A56EFC4277F25BFBEA873FC6FFCA2" authorityName="Meyer" authorityYear="1832" box="[721,934,783,805]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25BFBEA873FC6FFCA2" box="[721,934,783,805]" italics="true" pageId="4">Bottosaurus harlani</emphasis>
</taxonomicName>
frontal regions in dorsal view. A, TMM 40148-7, holotype,
<taxonomicName id="4C2A56EFC4277F25BC66A84EFE4CFCC4" authorityName="Cossette" authorityYear="2021" box="[265,389,813,835]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4277F25BC66A84EFE4CFCC4" box="[265,389,813,835]" italics="true" pageId="4">B. fustidens</emphasis>
</taxonomicName>
B, TMM 41366-16,
<taxonomicName id="4C2A56EFC4277F25BF37A84EFD1DFCC4" authorityName="Cossette" authorityYear="2021" box="[600,724,813,835]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4277F25BF37A84EFD1DFCC4" box="[600,724,813,835]" italics="true" pageId="4">B. fustidens</emphasis>
</taxonomicName>
C, NJSM 11265,
<taxonomicName id="4C2A56EFC4277F25BEE1A84EFC33FCC4" authorityName="Meyer" authorityYear="1832" box="[910,1018,813,835]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25BEE1A84EFC33FCC4" box="[910,1018,813,835]" italics="true" pageId="4">B. harlani</emphasis>
</taxonomicName>
. Abbreviations: f, frontal; pf, prefrontal. Scale bar = 5 cm.
</paragraph>
</caption>
<paragraph id="8B952D6CC4277F25BDCCAEB7FE24F9A0" blockId="4.[163,779,1492,1575]" pageId="4">
<emphasis id="B95EF17EC4277F25BDCCAEB7FE4FFA6D" box="[163,390,1492,1514]" italics="true" pageId="4">Referred specimens:</emphasis>
TMM 41366-16, mostly complete upper jaw, fragment of lower jaw, and partial elements of the posterior skull (
<figureCitation id="131131E9C4277F25BCF1AD72FE17F9A0" box="[414,478,1553,1575]" captionStart="Figure 7" captionStartId="11.[145,224,1264,1286]" captionTargetBox="[145,1425,195,1224]" captionTargetId="figure-282@11.[145,1425,195,1224]" captionTargetPageId="11" captionText="Figure 7. TMM 41366-16, Bottosaurus fustidens snout and posterior skull. AC, snout in dorsal (A), ventral (B) and right lateral (C) views. D, left posterolateral skull in dorsal view. E, dentary fragment in dorsal view. Abbreviations: d, dentary; f, frontal; j, jugal; l, lacrimal; mx, maxilla; mx#, numbered maxillary tooth; pf, prefrontal; pmx, premaxilla; q, quadrate; qj, quadratojugal; sq, squamosal. Scale bar = 5 cm." pageId="4">Fig. 7</figureCitation>
).
</paragraph>
<paragraph id="8B952D6CC4277F25BDCCAD33FE30F924" blockId="4.[163,778,1616,1699]" pageId="4">
<emphasis id="B95EF17EC4277F25BDCCAD33FEF3F9E2" box="[163,314,1616,1637]" italics="true" pageId="4">Occurrence:</emphasis>
Black Peaks Formation, Brewster County, TX,
<collectingCountry id="F33D6DFCC4277F25BC40AD0CFEACF903" box="[303,357,1647,1668]" name="United States of America" pageId="4">USA</collectingCountry>
(
<figureCitation id="131131E9C4277F25BC1BAD0CFE7DF903" box="[372,436,1647,1669]" captionStart="Figure 8" captionStartId="12.[163,245,762,784]" captionTargetBox="[321,1284,195,723]" captionTargetId="figure-24@12.[321,1284,195,723]" captionTargetPageId="12" captionText="Figure 8. Map of the USA showing the locations of Bottosaurus fustidens and Bottosaurus harlani fossil-bearing localities." pageId="4">Fig. 8</figureCitation>
); Palaeocene, Tiffanian North American Land Mammal Age.
</paragraph>
<paragraph id="8B952D6CC4277F25BDCCADAFFE39F898" blockId="4.[163,778,1740,1823]" pageId="4">
Etymology: From Latin
<emphasis id="B95EF17EC4277F25BCAFADAFFDC9F966" box="[448,512,1740,1761]" italics="true" pageId="4">fustis</emphasis>
, club, and
<emphasis id="B95EF17EC4277F25BFECADAFFD71F966" box="[643,696,1740,1761]" italics="true" pageId="4">dens</emphasis>
, tooth, to describe the robustly built, club-like mid-jaw and posterior teeth of the species.
</paragraph>
<paragraph id="8B952D6CC4277F25BDCCAC2AFA85FA95" blockId="4.[163,778,1864,1917]" lastBlockId="4.[827,1442,908,1298]" pageId="4">
<emphasis id="B95EF17EC4277F25BDCCAC2AFD89F8DA" box="[163,576,1864,1886]" italics="true" pageId="4">
Diagnosis:
<taxonomicName id="4C2A56EFC4277F25BC5DAC2AFD89F8DA" authorityName="Cossette" authorityYear="2021" box="[306,576,1864,1886]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="fustidens">Bottosaurus fustidens</taxonomicName>
</emphasis>
shares features with
<taxonomicName id="4C2A56EFC4277F25BD8BAC04FE17F8FB" authorityName="Meyer" authorityYear="1832" box="[228,478,1895,1916]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25BD8BAC04FE17F8FB" box="[228,478,1895,1916]" italics="true" pageId="4">Bottosaurus harlani</emphasis>
</taxonomicName>
but is differentiated by skull, mandibular and postcranial character states. A U-shaped depression similar to, but considerably shallower than that of
<taxonomicName id="4C2A56EFC4277F25B908A8A9FB20FC59" authorityName="Meyer" authorityYear="1832" box="[1127,1257,969,991]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25B908A8A9FB20FC59" box="[1127,1257,969,991]" italics="true" pageId="4">B. harlani</emphasis>
</taxonomicName>
, is present on the frontal at the point of the greatest mediolateral constriction between the orbits (
<figureCitation id="131131E9C4277F25B989AF64FAF9FB9B" box="[1254,1328,1031,1053]" captionStart="Figure 3" captionStartId="4.[163,242,783,805]" captionTargetBox="[323,1283,195,744]" captionTargetId="figure-481@4.[323,1283,195,744]" captionTargetPageId="4" captionText="Figure 3. Comparison of Bottosaurus fustidens and Bottosaurus harlani frontal regions in dorsal view. A, TMM 40148-7, holotype, B. fustidens B, TMM 41366-16, B. fustidens C, NJSM 11265, B. harlani. Abbreviations: f, frontal; pf, prefrontal. Scale bar = 5 cm." pageId="4">Fig. 3</figureCitation>
). Unlike
<taxonomicName id="4C2A56EFC4277F25BE54AF45FC67FBBD" authorityName="Meyer" authorityYear="1832" box="[827,942,1061,1083]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25BE54AF45FC67FBBD" box="[827,942,1061,1083]" italics="true" pageId="4">B. harlani</emphasis>
</taxonomicName>
, the splenial of
<taxonomicName id="4C2A56EFC4277F25B90EAF45FB2CFBBD" authorityName="Cossette" authorityYear="2021" box="[1121,1253,1061,1083]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="fustidens">
<emphasis id="B95EF17EC4277F25B90EAF45FB2CFBBD" box="[1121,1253,1061,1083]" italics="true" pageId="4">B. fustidens</emphasis>
</taxonomicName>
does not contact the mandibular symphysis, but is separated by a distance of two alveoli; its anteriormost projection passes ventral to the Meckelian groove (
<figureCitation id="131131E9C4277F25B872AFE2FAA9FB10" box="[1309,1376,1153,1175]" captionStart="Figure 9" captionStartId="13.[145,224,1817,1839]" captionTargetBox="[305,1265,1075,1778]" captionTargetId="figure-475@13.[305,1265,1075,1778]" captionTargetPageId="13" captionText="Figure 9. Comparison of Bottosaurus fustidens and Bottosaurus harlani splenials and mandibular symphyses in medial view. A, TMM 40148-7, holotype, B. fustidens. B, ANSP 9226, lectotype, B. harlani. Abbreviations: d, dentary; mg, Meckelian groove; ms, mandibular symphysis; s, splenial. Scale bar = 5 cm." pageId="4">Fig. 9</figureCitation>
). The mandibular tooth row posterior to the fifth alveolus is linear, in opposition to
<taxonomicName id="4C2A56EFC4277F25B930AFDCFB1DFB53" authorityName="Meyer" authorityYear="1832" box="[1119,1236,1215,1236]" class="Reptilia" family="Alligatoridae" genus="Bottosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="4" phylum="Chordata" rank="species" species="harlani">
<emphasis id="B95EF17EC4277F25B930AFDCFB1DFB53" box="[1119,1236,1215,1236]" italics="true" pageId="4">B. harlani</emphasis>
</taxonomicName>
. Dorsal vertebrae bear an anteroposteriorly oriented indentation along the ventral side of the vertebral body (
<figureCitation id="131131E9C4277F25B99FAF9FFAF5FA96" box="[1264,1340,1276,1298]" captionStart="Figure 10" captionStartId="14.[164,242,647,669]" captionTargetBox="[324,1284,196,607]" captionTargetId="figure-590@14.[324,1284,196,607]" captionTargetPageId="14" captionText="Figure 10. Comparison of Bottosaurus fustidens and Bottosaurus harlani dorsal vertebrae in ventral view. A, TMM 40148- 7, holotype, B. fustidens. B, NJSM 11265, B. harlani." pageId="4">Fig. 10</figureCitation>
).
</paragraph>
<paragraph id="8B952D6CC4277F25BE54AE58FADBFA6E" blockId="4.[827,1443,1339,1514]" pageId="4">
<emphasis id="B95EF17EC4277F25BE54AE58FC1CFAD7" box="[827,981,1339,1360]" italics="true" pageId="4">Description:</emphasis>
Morphology is described from
<specimenCount id="9D2CE6E5C4277F25B819AE58FC66FAE8" pageId="4" type="generic">two specimens</specimenCount>
, the
<typeStatus id="549193CEC4277F25BE8AAE3AFB8DFAE8" box="[997,1092,1369,1391]" pageId="4" type="holotype">holotype</typeStatus>
individual (TMM 40148-7) and a smaller referred individual (TMM 41366-16) from the same locality and horizon. Preservation of TMM 40148-7 is better than that of TMM 41366-16; the former is more complete and has clearer sutural contacts.
</paragraph>
</subSubSection>
</treatment>
</document>