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<document id="3DDD6A17D823F6ACBD6EDD4FD390774C" ID-DOI="10.1080/14772019.2022.2091959" ID-ISSN="1478-0941" ID-ZooBank=":pub:D3D3B15B-36FA-42EB-98AD-FAF369D" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1711485825671" checkinUser="felipe" docAuthor="Gracia, Carlos De, Correa-Metrio, Alex, Carvalho, Monica, Velez-Juarbe, Jorge, P ̆ rik, Tom ́ a ̆ s, , Carlos Jarami &amp; Kri, nd Jurgen" docDate="2022" docId="03CA87C46F218202FE92FA33FE7C26B2" docLanguage="en" docName="JSystPalaeontol.20.1.1-36.pdf" docOrigin="Journal of Systematic Palaeontology (2091959) 20 (1)" docSource="http://dx.doi.org/10.1080/14772019.2022.2091959" docStyle="DocumentStyle:6400638016C1F4AE0D0B6502E032D571.1:JSystPalaeontol.2016-.journal_article.closed" docStyleId="6400638016C1F4AE0D0B6502E032D571" docStyleName="JSystPalaeontol.2016-.journal_article.closed" docStyleVersion="1" docTitle="Morgula donosochagrense Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri 2022, sp. nov." docType="treatment" docVersion="1" lastPageNumber="13" masterDocId="FFF3FFBC6F2A820FFF99FF8CFF88256D" masterDocTitle="Towards a unifying systematic scheme of fossil and living billfishes (Teleostei, Istiophoridae)" masterLastPageNumber="36" masterPageNumber="1" pageNumber="11" updateTime="1711485829411" updateUser="felipe">
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<mods:title id="4F8C27CABA52C913F59769E7D7640FD8">Towards a unifying systematic scheme of fossil and living billfishes (Teleostei, Istiophoridae)</mods:title>
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<mods:namePart id="CBB83E2E53AAF597E9AD9C4BDAE893F1">Gracia, Carlos De</mods:namePart>
<mods:affiliation id="6ADB28356709E850D186C8DBAB795CEC">Department of Palaeontology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, Josef Holaubek Platz 2 (UZA II) / 2 A 323, 1090, Vienna, Austria; &amp; Vienna Doctoral School of Ecology and Evolution (VDSEE), University of Vienna, Vienna, Austria; &amp; Smithsonian Tropical Research Institute, Box 0843 - 03092, Balboa, Ancon, Panama, Republic of Panama;</mods:affiliation>
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<mods:namePart id="D81D66C10341F48BB8E5E4CE256971A4">Correa-Metrio, Alex</mods:namePart>
<mods:affiliation id="22B0253666C18E7B98DA8F30B1F7DA12">Centro de Geociencias, Universidad Nacional Autonoma de M ́ exico, Campus Juriquilla, Quer ́ etaro 76230, Mexico; &amp; Instituto de Geolog ́ ıa, Universidad Nacional Autonoma de M ́ exico, Ciudad Universitaria, 04510, M ́ exico, D. F., Mexico;</mods:affiliation>
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<mods:namePart id="6F5ADF120B64F53748F8EB0FF860625F">Carvalho, Monica</mods:namePart>
<mods:affiliation id="46C390A88F11AC0996551BE7E6F7B8BE">Smithsonian Tropical Research Institute, Box 0843 - 03092, Balboa, Ancon, Panama, Republic of Panama;</mods:affiliation>
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<mods:namePart id="FF87E5E23110635EFEA22262C7B01C49">Velez-Juarbe, Jorge</mods:namePart>
<mods:affiliation id="739357C17644620131B8A25591A859B5">Department of Mammalogy, Natural History Museum of Los Angeles Country, 900 Exposition Boulevard, Los Angeles, CA 90007, USA; &amp; Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA;</mods:affiliation>
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<mods:namePart id="A493B134833B9BD2F7DE71DFA2C904FE">P ̆ rik, Tom ́ a ̆ s</mods:namePart>
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<mods:namePart id="E2201C1D49DA9FF216A0C106C685B0F5">, Carlos Jarami</mods:namePart>
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<mods:namePart id="C590FA9FE0FC457546B768AD6BB35648">Kri, nd Jurgen</mods:namePart>
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<mods:title id="6C2B1C867EEA967EFA3B49AE6E265A10">Journal of Systematic Palaeontology</mods:title>
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<mods:part id="DB7BB6598E15D7AEA2AED8F7F6C9D8B8">
<mods:date id="B24502C3FA89084594F6159353645E7A">2022</mods:date>
<mods:detail id="2B149FB58A8E126D462EBA3FC014D521" type="series">
<mods:title id="6620E859EE9E095908BCBC825E09FE8E">2091959</mods:title>
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<mods:number id="11752B21179CD65B67BD656D1E385254">2022-08-19</mods:number>
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<mods:number id="0674D15ACE054E96CC511DEF2046B8E2">20</mods:number>
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<mods:number id="A0999089A25EC79AAF1187421D979C91">1</mods:number>
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<mods:url id="111859127797B86340F74DC3F5ABB58A">http://dx.doi.org/10.1080/14772019.2022.2091959</mods:url>
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<mods:classification id="DB2FE6F666E81D612D264C218874874C">journal article</mods:classification>
<mods:identifier id="7CD2428689CD6B470251A47CCE6AF098" type="DOI">10.1080/14772019.2022.2091959</mods:identifier>
<mods:identifier id="DB49EBCD0DFA024C3749314946BC27BB" type="ISSN">1478-0941</mods:identifier>
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<treatment id="03CA87C46F218202FE92FA33FE7C26B2" LSID="urn:lsid:plazi:treatment:03CA87C46F218202FE92FA33FE7C26B2" httpUri="http://treatment.plazi.org/id/03CA87C46F218202FE92FA33FE7C26B2" lastPageId="13" lastPageNumber="13" pageId="11" pageNumber="11">
<subSubSection id="C37965596F218204FE92FA33FDF52094" pageId="11" pageNumber="11" type="nomenclature">
<paragraph id="8BDC36D26F218204FE92FA33FD2920BB" blockId="11.[267,673,1471,1529]" box="[267,673,1471,1495]" pageId="11" pageNumber="11">
<heading id="D09481BE6F218204FE92FA33FD2920BB" box="[267,673,1471,1495]" centered="true" fontSize="10" level="2" pageId="11" pageNumber="11" reason="8">
<taxonomicName id="4C634D516F218204FE80FA33FDC920BB" authority="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri, 2022" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" box="[281,577,1471,1494]" family="Istiophoridae" genus="Morgula" kingdom="Animalia" order="Perciformes" pageId="11" pageNumber="11" phylum="Chordata" rank="species" species="donosochagrense" status="sp. nov.">
<emphasis id="B917EAC06F218204FE80FA33FDC920BB" bold="true" box="[281,577,1471,1494]" italics="true" pageId="11" pageNumber="11">Morgula donosochagrense</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A22457BB6F218204FDD3FA33FD2920BB" box="[586,673,1471,1494]" pageId="11" pageNumber="11" rank="species">sp. nov.</taxonomicNameLabel>
</heading>
</paragraph>
<paragraph id="8BDC36D26F218204FEB6FA6EFDF52094" blockId="11.[267,673,1471,1529]" box="[303,637,1505,1529]" pageId="11" pageNumber="11">
(Figs 5
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<collectionCode id="ED72AE176F218204FE05FA6EFE382094" box="[412,432,1506,1529]" pageId="11" pageNumber="11">C</collectionCode>
, G, H, 6
<collectionCode id="ED72AE176F218204FD88FA6EFDAE2094" box="[529,550,1506,1529]" pageId="11" pageNumber="11">A</collectionCode>
, B, 7
<collectionCode id="ED72AE176F218204FDFBFA6EFDFC2094" box="[610,628,1506,1529]" pageId="11" pageNumber="11">C</collectionCode>
)
</paragraph>
</subSubSection>
<subSubSection id="C37965596F218204FF06F9BEFA232438" pageId="11" pageNumber="11" type="diagnosis">
<paragraph id="8BDC36D26F218204FF06F9BEFC842220" blockId="11.[159,781,1586,1901]" pageId="11" pageNumber="11">
<emphasis id="B917EAC06F218204FF06F9BEFE9C2324" bold="true" box="[159,276,1586,1609]" pageId="11" pageNumber="11">Diagnosis.</emphasis>
The new species is characterized by the following combination of morphological traits (autapomorphic characters are indicated by an asterisk): slender rostrum along its entire length combined with a circular cross-section at 0.25
<collectionCode id="ED72AE176F218204FE16F938FE1723A6" box="[399,415,1716,1739]" pageId="11" pageNumber="11">L</collectionCode>
(
<figureCitation id="13582A576F218204FE2EF938FD9823A6" box="[439,528,1716,1739]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6B</figureCitation>
); premaxillae divided into two separate bones for about 60% of the total rostral length$; the internal canals circular (
<tableCitation id="C6E103696F218204FD1FF979FD572261" box="[646,735,1781,1804]" captionStart="Table 3" captionStartId="9.[159,218,187,208]" captionText="Table 3. Relationship between height of left nutrient canal at 0.5 L (H1) with the CA1/AR1 ratio (canal area / cross-section area at 0.5 L). Using H1 as size estimator, small nutrient canals range between 1.0 and 3.9 mm and big canals range between 4.0 and 8.5 mm. The ratio CA1/AR1 removes the size effect and small canals have values CA1/AR1 of 0.0020.019 and big canals of 0.020.07. In the score column b is big and s is small for the H1 and CA1/AR1 variables. For †Makaira purdyi variables at 0.25 L were used given the special morphology of this species. Ranges were defined after studying fossils and modern specimens together (Table S1)." pageId="11" pageNumber="11">Table 3</tableCitation>
) in ventral position (
<figureCitation id="13582A576F218204FEC2F899FE592241" box="[347,465,1813,1836]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A, B</figureCitation>
); rostrum elongated triangular in dorsal view$; nasals angularly extended laterally
</paragraph>
<paragraph id="8BDC36D26F218204FF06F8DAFEC32200" blockId="11.[159,781,1586,1901]" box="[159,331,1870,1901]" pageId="11" pageNumber="11">
(Figs 2
<collectionCode id="ED72AE176F218204FF72F8DAFF772200" box="[235,255,1878,1901]" pageId="11" pageNumber="11">A</collectionCode>
, 5
<collectionCode id="ED72AE176F218204FE82F8DAFEA62200" box="[283,302,1878,1901]" pageId="11" pageNumber="11">C</collectionCode>
)$.
</paragraph>
<paragraph id="8BDC36D26F218204FCD4FF31FA232438" blockId="11.[845,1467,189,341]" pageId="11" pageNumber="11">
<emphasis id="B917EAC06F218204FCD4FF31FB0825B9" bold="true" box="[845,1152,189,212]" pageId="11" pageNumber="11">Derivation of name.</emphasis>
The species name
<taxonomicName id="4C634D516F218204FCD4FF50FB842599" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" box="[845,1036,220,244]" family="Istiophoridae" genus="Morgula" kingdom="Animalia" order="Perciformes" pageId="11" pageNumber="11" phylum="Chordata" rank="species" species="donosochagrense">
<emphasis id="B917EAC06F218204FCCCFF51FB842599" box="[853,1036,221,244]" italics="true" pageId="11" pageNumber="11">donosochagrense</emphasis>
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is coined from a combination of the words Donoso and Chagres, the names of the two districts in which the outcrops of the Chagres Formation are located including parts of the natural reserve of Donoso.
</paragraph>
</subSubSection>
<subSubSection id="C37965596F218204FCD4FEE2FB7C271B" pageId="11" pageNumber="11" type="materials_examined">
<paragraph id="8BDC36D26F218204FCD4FEE2FBC224A8" blockId="11.[845,1467,365,453]" pageId="11" pageNumber="11">
<materialsCitation id="3B0B3C8F6F218204FCD4FEE2FBC224A8" collectionCode="A, C" pageId="11" pageNumber="11" specimenCount="1" typeStatus="holotype">
<emphasis id="B917EAC06F218204FCD4FEE2FC3324E8" bold="true" box="[845,955,366,389]" pageId="11" pageNumber="11">
<typeStatus id="54D888706F218204FCD4FEE2FC3F24E8" box="[845,951,366,389]" pageId="11" pageNumber="11" type="holotype">Holotype</typeStatus>
.
</emphasis>
MUPANSTRI31293, a nearly complete rostrum (Fig. 5
<collectionCode id="ED72AE176F218204FC4BFE02FC6F24C8" box="[978,999,398,421]" pageId="11" pageNumber="11">A</collectionCode>
<collectionCode id="ED72AE176F218204FC6DFE02FB8E24C8" box="[1012,1030,398,421]" pageId="11" pageNumber="11">C</collectionCode>
) with associated lower jaw lacking the predentary (Fig. 5GI).
</materialsCitation>
</paragraph>
<paragraph id="8BDC36D26F218204FCD4FE52FA322778" blockId="11.[844,1467,478,630]" pageId="11" pageNumber="11">
<emphasis id="B917EAC06F218204FCD4FE52FC512498" bold="true" box="[845,985,478,501]" pageId="11" pageNumber="11">Occurrence.</emphasis>
The specimen was collected in 2011 from a sea cliff exposure on the beach of the town of Pina
</paragraph>
<paragraph id="8BDC36D26F218204FCD4FD92FB7C271B" blockId="11.[844,1467,478,630]" pageId="11" pageNumber="11">
(
<figureCitation id="13582A576F218204FCCCFD92FC1F2758" box="[853,919,542,565]" captionStart="Figure 1" captionStartId="2.[159,226,950,971]" captionTargetBox="[159,1470,188,930]" captionTargetId="figure-453@2.[159,1470,188,933]" captionTargetPageId="2" captionText="Figure 1. AC, location map; stratigraphy and geological setting of the type locality (Pina, Chagres district, Costa Abajo, Colon Panama) after Collins et al. (1996)." pageId="11" pageNumber="11">Fig. 1</figureCitation>
), about 20 metres west of 9
<superScript id="7C169B9A6F218204FB4DFD96FB552746" attach="none" box="[1236,1245,538,555]" fontSize="7" pageId="11" pageNumber="11">ǫ</superScript>
9
<superScript id="7C169B9A6F218204FB75FD96FB792746" attach="none" box="[1260,1265,538,555]" fontSize="7" pageId="11" pageNumber="11">Ɩ</superScript>
16
<emphasis id="B917EAC06F218204FA94FD93FA9F275B" bold="true" box="[1293,1303,543,566]" italics="true" pageId="11" pageNumber="11">ʺ</emphasis>
N; 80
<superScript id="7C169B9A6F218204FACEFD96FAE82746" attach="none" box="[1367,1376,538,555]" fontSize="7" pageId="11" pageNumber="11">ǫ</superScript>
2
<superScript id="7C169B9A6F218204FAF6FD96FAFC2746" attach="none" box="[1391,1396,538,555]" fontSize="7" pageId="11" pageNumber="11">Ɩ</superScript>
71
<emphasis id="B917EAC06F218204FA09FD93FA12275B" bold="true" box="[1424,1434,543,566]" italics="true" pageId="11" pageNumber="11">ʺ</emphasis>
W. The fossiliferous unit is ~
<quantity id="4C9B9B376F218204FBE9FDB3FB15273B" box="[1136,1181,575,598]" metricMagnitude="0" metricUnit="m" metricValue="5.0" pageId="11" pageNumber="11" unit="m" value="5.0">5 m</quantity>
above a massive bioturbated layer that is exposed at tide level.
</paragraph>
</subSubSection>
<subSubSection id="C37965596F218202FCD5FD03FE7C26B2" lastPageId="13" lastPageNumber="13" pageId="11" pageNumber="11" type="discussion">
<paragraph id="8BDC36D26F218204FCD5FD03FBE8262A" blockId="11.[844,1467,655,1901]" pageId="11" pageNumber="11">
<emphasis id="B917EAC06F218204FCD5FD03FC3327CB" bold="true" box="[844,955,655,678]" pageId="11" pageNumber="11">Remarks.</emphasis>
The specimen was found in a weathered block above the tide zone. The surface of the fossil consequently is worn (Fig. 2
<collectionCode id="ED72AE176F218204FBFFFD43FBF2278B" box="[1126,1146,719,742]" pageId="11" pageNumber="11">A</collectionCode>
<collectionCode id="ED72AE176F218204FB11FD43FB12278B" box="[1160,1178,719,742]" pageId="11" pageNumber="11">C</collectionCode>
) and alveoli for denticles were not recognized. The specimen includes an associated lower jaw (Fig. 2GI) that was found in the same block during preparation.
</paragraph>
<paragraph id="8BDC36D26F218204FCFEFCDCFB272200" blockId="11.[844,1467,655,1901]" pageId="11" pageNumber="11">
The rostrum of †
<taxonomicName id="4C634D516F218204FBA8FCDCFAD0260A" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" box="[1073,1368,848,871]" family="Istiophoridae" genus="Morgula" kingdom="Animalia" order="Perciformes" pageId="11" pageNumber="11" phylum="Chordata" rank="species" species="donosochagrense">
<emphasis id="B917EAC06F218204FBA8FCDCFAD0260A" box="[1073,1368,848,871]" italics="true" pageId="11" pageNumber="11">Morgula donosochagrense</emphasis>
</taxonomicName>
gen. et. sp. nov. is nearly complete and consists of prenasals, premaxillaries and a complete left nasal (Fig. 5
<collectionCode id="ED72AE176F218204FA19FC1CFA1D26CA" box="[1408,1429,912,935]" pageId="11" pageNumber="11">A</collectionCode>
<collectionCode id="ED72AE176F218204FA3BFC1CFA3E26CA" box="[1442,1462,912,935]" pageId="11" pageNumber="11">C</collectionCode>
, G I). The posterior edge of the left nasal is in the same transverse plane as the lateral ethmoid (Fig. 2), which allows a comparison with other taxa (
<bibRefCitation id="EFF24B236F218204FAC2FC7DFC062145" author="Fierstine, H. L." pageId="11" pageNumber="11" pagination="430 - 437" refId="ref25205" refString="Fierstine, H. L. 1999 a. Makaira sp., cf. M. nigricans Lac´ep`ede, 1802 (Teleostei: Perciformes: Istiophoridae) from the late Miocene, Panama, and its probable use of the Panama seaway. Journal of Vertebrate Paleontology, 19, 430 - 437. doi: 10.1080 / 02724634.1999.10011156" type="journal article" year="1999">Fierstine 1999a</bibRefCitation>
). The overall preserved length of the rostrum is
<quantity id="4C9B9B376F218204FCD5FBBDFC2A2125" box="[844,930,1073,1096]" metricMagnitude="-1" metricUnit="m" metricValue="6.11" pageId="11" pageNumber="11" unit="mm" value="611.0">611 mm</quantity>
and only
<emphasis id="B917EAC06F218204FB8CFBBDFBA92125" box="[1045,1057,1073,1096]" italics="true" pageId="11" pageNumber="11">c</emphasis>
.
<quantity id="4C9B9B376F218204FBA8FBBDFBF12125" box="[1073,1145,1073,1096]" metricMagnitude="-2" metricUnit="m" metricValue="1.3" pageId="11" pageNumber="11" unit="mm" value="13.0">13 mm</quantity>
of the distal tip are missing. The estimated total bill length therefore is
<quantity id="4C9B9B376F218204FAFDFBDDFA322105" box="[1380,1466,1105,1128]" metricMagnitude="-1" metricUnit="m" metricValue="6.24" pageId="11" pageNumber="11" unit="mm" value="624.0">624 mm</quantity>
(Supplemental material 1). The rostrum s cross-section is oval at 0.5
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(
<quantity id="4C9B9B376F218204FB8AFB1EFBE121C4" box="[1043,1129,1170,1193]" metricMagnitude="-1" metricUnit="m" metricValue="3.07" pageId="11" pageNumber="11" unit="mm" value="307.0">307 mm</quantity>
from the tip) but circular at 0.25
<collectionCode id="ED72AE176F218204FC19FB3EFC1821A4" box="[896,912,1202,1225]" pageId="11" pageNumber="11">L</collectionCode>
(
<quantity id="4C9B9B376F218204FC3AFB3EFC7121A4" box="[931,1017,1202,1225]" metricMagnitude="-1" metricUnit="m" metricValue="1.55" pageId="11" pageNumber="11" unit="mm" value="155.0">155 mm</quantity>
from the tip [
<figureCitation id="13582A576F218204FB38FB3EFA9321A4" box="[1185,1307,1202,1225]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A, B</figureCitation>
]; see the outrange ratio D2/W
<quantity id="4C9B9B376F218204FB95FB5EFBBF2184" box="[1036,1079,1234,1257]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="11" pageNumber="11" unit="in" value="2.0">2 in</quantity>
<tableCitation id="C6E103696F218204FBD9FB5EFB1A2184" box="[1088,1170,1234,1257]" captionStart="Table 1" captionStartId="7.[133,192,475,496]" captionText="Table 1. Mean (x), observed range, and number of specimens examined (n) for nine rostral and one dentary ratios of eight Istiophoridae billfish compared with three fossil species. Abbreviations of the ratio are explained in the methodology and in the legends of Figures 3, 5 and 6. Values presented here for marlins were obtained after data imputation. The variable DD1/D1 for K. audax and all data from spearfishes (Tetrapturus ssp.) were taken from Fierstine (2001). NA non-available data, $ indicates out of range values if compared with modern species." pageId="11" pageNumber="11">Table 1</tableCitation>
). The distal tip has a slen- der outline (Figs 5
<collectionCode id="ED72AE176F218204FB85FB7EFBB92064" box="[1052,1073,1266,1289]" pageId="11" pageNumber="11">A</collectionCode>
<collectionCode id="ED72AE176F218204FBA7FB7EFBD82064" box="[1086,1104,1266,1289]" pageId="11" pageNumber="11">C</collectionCode>
, Supplemental material S1B), is acute and dorsoventrally compressed. The premaxillaries start to separate at
<quantity id="4C9B9B376F218204FBBCFABFFBF22027" box="[1061,1146,1331,1354]" metricMagnitude="-1" metricUnit="m" metricValue="2.48" pageId="11" pageNumber="11" unit="mm" value="248.0">248 mm</quantity>
(
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) measured from the distal tip (Fig. 5
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). The
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distance represents approximately 30% of the total rostrum length and coincides with the point where the ventral canal starts to be visible in cross-section at 0.5
<collectionCode id="ED72AE176F218204FB39FA38FB3820A6" box="[1184,1200,1460,1483]" pageId="11" pageNumber="11">L</collectionCode>
(
<figureCitation id="13582A576F218204FB5FFA38FA9620A6" box="[1222,1310,1460,1483]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A</figureCitation>
). The ventral aperture (
<collectionCode id="ED72AE176F218204FC25FA58FC462086" box="[956,974,1492,1515]" pageId="11" pageNumber="11">L</collectionCode>
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) represents 60% of the rostral length (
<figureCitation id="13582A576F218204FC38FA78FC7C2366" box="[929,1012,1524,1547]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A</figureCitation>
). In dorsal view, the nasals are angularly oriented posterolaterally (
<figureCitation id="13582A576F218204FBF7F998FB4C2346" box="[1134,1220,1556,1579]" captionStart="Figure 7" captionStartId="15.[159,226,680,701]" captionTargetBox="[169,1457,192,661]" captionTargetId="figure-417@15.[168,1457,188,663]" captionTargetPageId="15" captionText="Figure 7. AL, comparison among selected fossils and extant marlin species with emphasis on the rostral morphology and its crosssections at 0.5 and 0.25L; A, reconstruction of the rostrum of †Prototetrapturus courcelli gen. et. comb. nov. (MNHN P250); B, reconstruction of the rostrum of †Spathochoira calvertense gen. et. comb. nov. (USNM 9344); C, †Morgula donosochagrense gen. et sp. nov. (MUPANSTRI31293); D, †Makaira colonense sp. nov. (MUPANSTRI31292); E, reconstruction of the rostrum of Makaira panamense (USNM 181710); F, reconstruction of the rostrum of †Makaira belgica (MAUL 917/1) from the late Miocene of Italy; G, †Makaira fierstini sp. nov. (USNM 358534); H, Makaira nigricans (USNM 196019); I, Istiompax indica (LACM 25509); J, Kajikia albida (USNM 360507); K, Kajikia audax (USNM 372777); L, Istiophorus platypterus (MNHN A-9463). The white circle indicates the point where fused prenasals start to divide in two separate bones, if this point is more distal, the narial cavity is longer and rostrum is more open. Arrows indicates where premaxillaries protrude from prenasals in Makaira ssp., the outer projection of the prenasals is represented by an angle symbol. Cross-sections at 0.5 and 0.25L: blue is oval shape, light blue is oval shape compressed laterally (borders could be more angular instead of smooth and prenasals are embedded into premaxillaries; see Fig. 9H), green is round, yellow is oval highly depressed (almost plane), red represents other shapes (see Fig. 9A, J). Pink water mark indicates the reconstructed sections. Different rostrums are labelled according the observed or inferred feeding strategy. Thrusting refers to rostrums used mostly for direct impacts and specific refers to rostrums used in multidirectional, lateral or complex feeding strategies. Scale bars = 10 cm." pageId="11" pageNumber="11">Fig. 7C</figureCitation>
). The internal canals are large, ventrally located, centrally aligned, and with a circular cross-section (
<tableCitation id="C6E103696F218204FBC7F9D9FB362301" box="[1118,1214,1621,1644]" captionStart="Table 3" captionStartId="9.[159,218,187,208]" captionText="Table 3. Relationship between height of left nutrient canal at 0.5 L (H1) with the CA1/AR1 ratio (canal area / cross-section area at 0.5 L). Using H1 as size estimator, small nutrient canals range between 1.0 and 3.9 mm and big canals range between 4.0 and 8.5 mm. The ratio CA1/AR1 removes the size effect and small canals have values CA1/AR1 of 0.0020.019 and big canals of 0.020.07. In the score column b is big and s is small for the H1 and CA1/AR1 variables. For †Makaira purdyi variables at 0.25 L were used given the special morphology of this species. Ranges were defined after studying fossils and modern specimens together (Table S1)." pageId="11" pageNumber="11">Table 3</tableCitation>
;
<figureCitation id="13582A576F218204FB45F9D9FAE42301" box="[1244,1388,1621,1644]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A, B</figureCitation>
). The rounded internal canals that are ventrally placed are autapomorphies of †
<taxonomicName id="4C634D516F218204FBAEF919FAD623C1" authority="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri, 2022" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" box="[1079,1374,1685,1708]" family="Istiophoridae" genus="Morgula" kingdom="Animalia" order="Perciformes" pageId="11" pageNumber="11" phylum="Chordata" rank="species" species="donosochagrense" status="gen. et sp. nov.">
<emphasis id="B917EAC06F218204FBAEF919FAD623C1" box="[1079,1374,1685,1708]" italics="true" pageId="11" pageNumber="11">Morgula donosochagrense</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A22457BB6F218204FAF5F919FC2E23A1" pageId="11" pageNumber="11" rank="species">gen. et sp. nov.</taxonomicNameLabel>
(
<figureCitation id="13582A576F218204FC23F939FBBD23A1" box="[954,1077,1717,1740]" captionStart="Figure 6" captionStartId="14.[159,226,1365,1386]" captionTargetBox="[159,1469,205,1348]" captionTargetId="figure-217@14.[159,1489,188,1348]" captionTargetPageId="14" captionText="Figure 6. A, B, †Morgula donosochagrense gen et sp. nov. (MUPANSTRI31293); A, cross-section at 0.5L; B, cross-section at 0.25L. C, D, †Makaira colonense sp. nov. (MUPANSTRI39292); C, cross-section at 0.5L; D, cross-section at 0.25 L. Abbreviation: PN, prenasal. Scale bars = 1 cm." pageId="11" pageNumber="11">Fig. 6A, B</figureCitation>
), as is the triangular shape of the rostrum and the nasal bones that are angularly extended laterally in dorsal view (Figs 5
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<collectionCode id="ED72AE176F218204FB4EF97AFB612260" box="[1239,1257,1782,1805]" pageId="11" pageNumber="11">C</collectionCode>
, 7
<collectionCode id="ED72AE176F218204FA93F97AFA942260" box="[1290,1308,1782,1805]" pageId="11" pageNumber="11">C</collectionCode>
). Other istiophorids have oval internal canals in a central or dorsal position (see
<figureCitation id="13582A576F218204FC69F8BAFBB02220" box="[1008,1080,1846,1869]" captionStart="Figure 9" captionStartId="19.[159,226,1672,1693]" captionTargetBox="[165,779,192,1654]" captionTargetId="figure-292@19.[161,779,188,1654]" captionTargetPageId="19" captionText="Figure 9. AE, Cross-sections of fossil istiophorids. †Prototetrapturus courcelli gen. et. comb. nov. (MNHN P250): A, cross-section at the proximal end of the rostrum; B, crosssection at 92 mm measured from its proximal end; C, crosssection at 139 mm measured from its proximal end; D, crosssection at 200mm measured from its proximal end; E, crosssection at 260 mm measured from its proximal end, F, †Makaira purdyi (USNM 481933), cross-section at 0.25 L distance. G, †Makaira panamense (USNM 481933), cross-" pageId="11" pageNumber="11">Fig. 9</figureCitation>
for examples and Supplemental material Fig. S2 for definitions).
</paragraph>
<paragraph id="8BDC36D26F278202FF06FF31FE7C26B2" blockId="13.[159,781,189,991]" pageId="13" pageNumber="13">
<emphasis id="B917EAC06F278202FF06FF31FEAC25B9" bold="true" box="[159,292,189,212]" pageId="13" pageNumber="13">Lower jaw.</emphasis>
The associated lower jaw is represented by partial right and left dentaries (Fig. 5GI). In the articulation of the dentaries, two variables (DAD and
<collectionCode id="ED72AE176F278202FD4FFF72FC8E2478" box="[726,774,254,277]" pageId="13" pageNumber="13">DJL</collectionCode>
) were studied and converted into a ratio (
<figureCitation id="13582A576F278202FDEBFE92FD402458" box="[626,712,286,309]" captionStart="Figure 3" captionStartId="6.[159,226,1306,1327]" captionTargetBox="[167,772,198,1285]" captionTargetId="figure-507@6.[164,776,188,1285]" captionTargetPageId="6" captionText="Figure 3. A, B, characters studied with emphasis on two regions of the rostrum: 0.5L, half of the distance between the distal tip and the orbital margin of the lateral ethmoid bone and 0.25L, one-fourth the distance between the distal tip and the orbital margin of the lateral ethmoid bone; A, cross-section view at 0.5L; B, cross-section view at 0.25 L; C, left dentary on medial view of the interdentary joint. Character abbreviations: depth (D1, D2) and width (W1, W2) of the rostrum, height (H1, H2) and width (N1, N2) of the left internal canal (as seen in cross-section), intercanal distance (IC1, IC2) and distance of the internal canal from dorsal surface (DD1, DD2). Depth from the anterior-most denticles perpendicular to the ventral margin of the dentary (not to the distal margin, which often has projection) (DAD), and length of the intermediary joint from the mandibular foramen to the anterior-most denticles (DJL). Mandibular foramen = mf. Modified from Fierstine &amp; Voigt (1996) and Fierstine (2001)." pageId="13" pageNumber="13">Fig. 3C</figureCitation>
). The ratio DAD/DJL for †
<taxonomicName id="4C634D516F278202FE15FEB3FD38243B" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" box="[396,688,319,342]" family="Istiophoridae" genus="Morgula" kingdom="Animalia" order="Perciformes" pageId="13" pageNumber="13" phylum="Chordata" rank="species" species="donosochagrense">
<emphasis id="B917EAC06F278202FE15FEB3FD38243B" box="[396,688,319,342]" italics="true" pageId="13" pageNumber="13">Morgula donosochagrense</emphasis>
</taxonomicName>
gen. et. sp. nov. is 0.52 (
<tableCitation id="C6E103696F278202FEF2FED3FE4A241B" box="[363,450,351,374]" captionStart="Table 1" captionStartId="7.[133,192,475,496]" captionText="Table 1. Mean (x), observed range, and number of specimens examined (n) for nine rostral and one dentary ratios of eight Istiophoridae billfish compared with three fossil species. Abbreviations of the ratio are explained in the methodology and in the legends of Figures 3, 5 and 6. Values presented here for marlins were obtained after data imputation. The variable DD1/D1 for K. audax and all data from spearfishes (Tetrapturus ssp.) were taken from Fierstine (2001). NA non-available data, $ indicates out of range values if compared with modern species." pageId="13" pageNumber="13">Table 1</tableCitation>
) and this value represents a shape that is more rectangular than quadratic distally where both dentaries join (
<figureCitation id="13582A576F278202FE46FE2CFDB324DA" box="[479,571,416,439]" captionStart="Figure 3" captionStartId="6.[159,226,1306,1327]" captionTargetBox="[167,772,198,1285]" captionTargetId="figure-507@6.[164,776,188,1285]" captionTargetPageId="6" captionText="Figure 3. A, B, characters studied with emphasis on two regions of the rostrum: 0.5L, half of the distance between the distal tip and the orbital margin of the lateral ethmoid bone and 0.25L, one-fourth the distance between the distal tip and the orbital margin of the lateral ethmoid bone; A, cross-section view at 0.5L; B, cross-section view at 0.25 L; C, left dentary on medial view of the interdentary joint. Character abbreviations: depth (D1, D2) and width (W1, W2) of the rostrum, height (H1, H2) and width (N1, N2) of the left internal canal (as seen in cross-section), intercanal distance (IC1, IC2) and distance of the internal canal from dorsal surface (DD1, DD2). Depth from the anterior-most denticles perpendicular to the ventral margin of the dentary (not to the distal margin, which often has projection) (DAD), and length of the intermediary joint from the mandibular foramen to the anterior-most denticles (DJL). Mandibular foramen = mf. Modified from Fierstine &amp; Voigt (1996) and Fierstine (2001)." pageId="13" pageNumber="13">Figs 3C</figureCitation>
, 5I). The general appearance of the dentaries is slender with a height at the interdentary joint of
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(
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distance) and of
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at the level of the articular socket for the head of the quadrate. The right dentary is nearly complete with its proximal part missing ventrally for half of its length (Fig. 5G), while its dorsal surface is completely preserved (Fig. 5H). The left dentary is partially preserved but broken in its middle part at the location of the articular joint (Fig. 5G, H). In dorsal view, the dentary is covered by denticles from its most distal part to about the level of the articular socket (Fig. 5H). There is no further evidence of denticles from this point to the most proximal end. The surface that is covered with denticles presents circular structures of
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that are possibly alveoli for larger teeth (Fig. 5H). There are one and three of these structures visible on the right and left dentary, respectively (Fig. 5H).
</paragraph>
</subSubSection>
</treatment>
</document>