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<document id="29102819849886332122BC01DA898B3B" ID-DOI="10.1080/14772019.2022.2091959" ID-ISSN="1478-0941" ID-Zenodo-Dep="10979515" ID-ZooBank=":pub:D3D3B15B-36FA-42EB-98AD-FAF369D" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="diego" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="diego" IM.taxonomicNames_approvedBy="diego" IM.treatments_approvedBy="diego" 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="03CA87C46F3E8219FC19FE93FD9C27D8" 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="Prototetrapturus courcelli Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri 2022, comb. nov." docType="treatment" docVersion="4" lastPageNumber="20" masterDocId="FFF3FFBC6F2A820FFF99FF8CFF88256D" masterDocTitle="Towards a unifying systematic scheme of fossil and living billfishes (Teleostei, Istiophoridae)" masterLastPageNumber="36" masterPageNumber="1" pageNumber="20" updateTime="1713275199640" updateUser="ExternalLinkService">
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<mods:title id="CD183256AB5EBB835BA075E102E0EF62">Towards a unifying systematic scheme of fossil and living billfishes (Teleostei, Istiophoridae)</mods:title>
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<mods:namePart id="840296858A1BBD75DC2C7CB2BC676A4F">Gracia, Carlos De</mods:namePart>
<mods:affiliation id="FCA02F39285236BA992DE267079374A9">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="E2B15C18E671DFC1FECA2BC516FB2009">Correa-Metrio, Alex</mods:namePart>
<mods:affiliation id="317EA47951E85148C74FBB712B0E6EF3">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:affiliation id="739810F42E2EB5A440BFD3D20711EC78">Smithsonian Tropical Research Institute, Box 0843 - 03092, Balboa, Ancon, Panama, Republic of Panama;</mods:affiliation>
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<mods:namePart id="181AAEFE39A0E6910DBE8DA7D4E9E190">Velez-Juarbe, Jorge</mods:namePart>
<mods:affiliation id="C4F414120A72DF9320447B61BBA527C9">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="ECD9589C3CDFBDA1AF6C8E09E52A1603">P ̆ rik, Tom ́ a ̆ s</mods:namePart>
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<mods:namePart id="1DF95110293CCA7E818D54218B7C7F50">, Carlos Jarami</mods:namePart>
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<mods:date id="50D7A457AE15E6C22985B3E070B90E34">2022</mods:date>
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<paragraph id="8BDC36D26F3E821BFC19FE93FB492435" blockId="20.[864,1447,287,378]" pageId="20" pageNumber="20">
<taxonomicName id="4C634D516F3E821BFC14FEACFA0F245A" authority="(Arambourg, 1927)" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" baseAuthorityName="Arambourg" baseAuthorityYear="1927" box="[909,1415,288,311]" family="Istiophoridae" genus="Prototetrapturus" kingdom="Animalia" order="Perciformes" pageId="20" pageNumber="20" phylum="Chordata" rank="species" species="courcelli" status="comb. nov.">
<emphasis id="B917EAC06F3E821BFC14FEACFB24245A" bold="true" box="[909,1196,288,311]" italics="true" pageId="20" pageNumber="20">Prototetrapturus courcelli</emphasis>
(
<bibRefCitation id="EFF24B236F3E821BFB24FEACFAF6245A" author="Arambourg, C." box="[1213,1406,288,311]" pageId="20" pageNumber="20" pagination="1 - 298" refId="ref23620" refString="Arambourg, C. 1927. Les poissons fossiles d' Oran. Mat´eriaux pour la Carte g´eologiques de l' Alg´erie, S´erie 1, Pal´eontologie, 6, 1 - 298." type="journal article" year="1927">Arambourg, 1927</bibRefCitation>
)
</taxonomicName>
<taxonomicNameLabel id="A22457BB6F3E821BFBDEFECDFB492435" box="[1095,1217,321,344]" pageId="20" pageNumber="20" rank="species">comb. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="C37965596F3E821BFCF9FEEFFA2F2417" box="[864,1447,355,378]" pageId="20" pageNumber="20" type="description">
<paragraph id="8BDC36D26F3E821BFCF9FEEFFA2F2417" blockId="20.[864,1447,287,378]" box="[864,1447,355,378]" pageId="20" pageNumber="20">
(
<figureCitation id="13582A576F3E821BFCF0FEEFFC492417" box="[873,961,355,378]" 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="20" pageNumber="20">Figs 7B</figureCitation>
,
<figureCitation id="13582A576F3E821BFC56FEEFFC662417" box="[975,1006,355,378]" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="20" pageNumber="20">8C</figureCitation>
,
<figureCitation id="13582A576F3E821BFC67FEEFFBCB2417" box="[1022,1091,355,378]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="20" pageNumber="20">9AE</figureCitation>
, Supplemental material
<figureCitation id="13582A576F3E821BFAD7FEEFFA152417" box="[1358,1437,355,378]" captionStart="Figure 4" captionStartId="10.[159,226,1174,1195]" captionTargetBox="[159,1469,191,1149]" captionTargetId="figure-227@10.[159,1469,188,1157]" captionTargetPageId="10" captionText="Figure 4. Maximum likelihood constrained topology based on Williams et al. (2018) for istiophorid billfishes. Colours indicate different clades that are correlated with different types of rostra and behaviours based on extant species. The definition for the rostra shape is detailed in the character matrix (Supplemental material Table S2). The skull figures in ventral view show the morphology of the neurocranium, vomer and parasphenoid in different clades. The colours for skull bones are explained in Figure 13. The rostrum and drawings on the button show how each type of rostra correlated with a different feeding behaviours based on observations in extant species. The silhouettes of the three main types of rostra described are compared with drawings of different types of swords for better understanding of the rostral morphology. Green is extremely thin rostra similar to a fencing sword; blue illustrates robust rostrums compared with a gladius hispaniensis or Mainz-type swords, as shown in Bishop (2016, pp. 1821); red shows slender rostrums compared with a Viking sword (see Oakeshott [1960] for examples of this type of sword) and orange indicates the same rostrum type as red. Colours also show the convergence of different rostral, vomer and parasphenoid shapes in our topology. The Tetrapturus clade (orange) includes species with slender rostra similar to the Gracilorosta clade (red) but no information about the feeding behaviour in spearfishes is available." pageId="20" pageNumber="20">Fig. S4</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C37965596F3E821BFCD3FE2FFA242756" pageId="20" pageNumber="20" type="reference_group">
<paragraph id="8BDC36D26F3E821BFCD3FE2FFC5B24B6" blockId="20.[842,1466,419,572]" pageId="20" pageNumber="20">
1927 †
<taxonomicName id="4C634D516F3E821BFC0FFE2FFAA724D7" authority="Arambourg" authorityName="Arambourg" authorityYear="1927" box="[918,1327,419,442]" family="Xiphiidae" genus="Xiphiorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perciformes" pageId="20" pageNumber="20" phylum="Chordata" rank="species" species="courcelli">
<emphasis id="B917EAC06F3E821BFC0FFE2FFB2F24D7" box="[918,1191,419,442]" italics="true" pageId="20" pageNumber="20">Xiphiorhynchus courcelli</emphasis>
Arambourg
</taxonomicName>
: 173, pl. 36/ 6, fig. 40.
</paragraph>
<paragraph id="8BDC36D26F3E821BFCD3FE68FA162496" blockId="20.[842,1466,419,572]" box="[842,1438,483,507]" pageId="20" pageNumber="20">
1978 †
<taxonomicName id="4C634D516F3E821BFC0FFE68FA982496" authority="(Arambourg)" baseAuthorityName="Arambourg" box="[918,1296,484,507]" class="Mammalia" family="Istiophoridae" genus="Istiophorus" kingdom="Animalia" order="Perciformes" pageId="20" pageNumber="20" phylum="Chordata" rank="species" species="courcelli">
<emphasis id="B917EAC06F3E821BFC0FFE68FBFF2496" box="[918,1143,484,507]" italics="true" pageId="20" pageNumber="20">Istiophorus courcelli</emphasis>
(Arambourg)
</taxonomicName>
;
<bibCitation id="5E493EBE6F3E821BFA84FE68FA162496" box="[1309,1438,484,507]" page="8" pageId="20" pageNumber="20" score="1.97023359760351E-6" volumeTitle="Fierstine" year="1987">Fierstine: 8.</bibCitation>
</paragraph>
<paragraph id="8BDC36D26F3E821BFCD3FD88FA242756" blockId="20.[842,1466,419,572]" pageId="20" pageNumber="20">
1987 †
<taxonomicName id="4C634D516F3E821BFC06FD88FA852776" authority="(Arambourg)" baseAuthorityName="Arambourg" box="[927,1293,516,539]" family="Istiophoridae" genus="Makaira" kingdom="Animalia" order="Perciformes" pageId="20" pageNumber="20" phylum="Chordata" rank="species" species="courcelli">
<emphasis id="B917EAC06F3E821BFC06FD88FBE32776" box="[927,1131,516,539]" italics="true" pageId="20" pageNumber="20">Makaira courcelli</emphasis>
(Arambourg)
</taxonomicName>
; Schultz: 120, 140, 148, 172, tabs. 2, 3, figs 6A, 7C, 8AE, S1, S4.
</paragraph>
</subSubSection>
<subSubSection id="C37965596F3E821BFCD5FDD8FC2426FD" pageId="20" pageNumber="20" type="diagnosis">
<paragraph id="8BDC36D26F3E821BFCD5FDD8FC2426FD" blockId="20.[844,1467,596,912]" pageId="20" pageNumber="20">
<emphasis id="B917EAC06F3E821BFCD5FDD8FC492706" bold="true" box="[844,961,596,619]" pageId="20" pageNumber="20">Diagnosis.</emphasis>
<taxonomicName id="4C634D516F3E821BFC79FDD8FA8C2706" authorityName="Gracia &amp; Correa-Metrio &amp; Carvalho &amp; Velez-Juarbe &amp; P̆rik &amp; &amp; Kri" authorityYear="2022" baseAuthorityName="Arambourg" baseAuthorityYear="1927" box="[992,1284,596,619]" family="Istiophoridae" genus="Prototetrapturus" kingdom="Animalia" order="Perciformes" pageId="20" pageNumber="20" phylum="Chordata" rank="species" species="courcelli" status="comb. nov.">
<emphasis id="B917EAC06F3E821BFC79FDD8FA8C2706" box="[992,1284,596,619]" italics="true" pageId="20" pageNumber="20">Prototetrapturus courcelli</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A22457BB6F3E821BFA8CFDD8FA102706" box="[1301,1432,596,619]" pageId="20" pageNumber="20" rank="species">comb. nov.</taxonomicNameLabel>
is differentiated from all other istiophorids in the following combination of morphological traits (an asterisk indicates autapomorphic traits): very slender rostrum with acute distal tip; an oval to angularly obtuse cross-section$; two additional, small and dorsolaterally positioned internal canals in dorsal position and laterally distributed$; denticles on the lateral sides that are distinctly large, conical and pointed$ (Supplemental material
<figureCitation id="13582A576F3E821BFCD5FCF5FC1426FD" box="[844,924,889,912]" captionStart="Figure 4" captionStartId="10.[159,226,1174,1195]" captionTargetBox="[159,1469,191,1149]" captionTargetId="figure-227@10.[159,1469,188,1157]" captionTargetPageId="10" captionText="Figure 4. Maximum likelihood constrained topology based on Williams et al. (2018) for istiophorid billfishes. Colours indicate different clades that are correlated with different types of rostra and behaviours based on extant species. The definition for the rostra shape is detailed in the character matrix (Supplemental material Table S2). The skull figures in ventral view show the morphology of the neurocranium, vomer and parasphenoid in different clades. The colours for skull bones are explained in Figure 13. The rostrum and drawings on the button show how each type of rostra correlated with a different feeding behaviours based on observations in extant species. The silhouettes of the three main types of rostra described are compared with drawings of different types of swords for better understanding of the rostral morphology. Green is extremely thin rostra similar to a fencing sword; blue illustrates robust rostrums compared with a gladius hispaniensis or Mainz-type swords, as shown in Bishop (2016, pp. 1821); red shows slender rostrums compared with a Viking sword (see Oakeshott [1960] for examples of this type of sword) and orange indicates the same rostrum type as red. Colours also show the convergence of different rostral, vomer and parasphenoid shapes in our topology. The Tetrapturus clade (orange) includes species with slender rostra similar to the Gracilorosta clade (red) but no information about the feeding behaviour in spearfishes is available." pageId="20" pageNumber="20">Fig. S4</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37965596F3E821BFCD5FC25FBB3211F" pageId="20" pageNumber="20" type="materials_examined">
<paragraph id="8BDC36D26F3E821BFCD5FC25FBC3268C" blockId="20.[844,1467,937,993]" pageId="20" pageNumber="20">
<materialsCitation id="3B0B3C8F6F3E821BFCD5FC25FBC3268C" collectionCode="MNHNP" pageId="20" pageNumber="20" specimenCode="MNHNP 250" specimenCount="1" typeStatus="holotype">
<emphasis id="B917EAC06F3E821BFCD5FC25FC3326AD" bold="true" box="[844,955,937,960]" pageId="20" pageNumber="20">
<typeStatus id="54D888706F3E821BFCD5FC25FC3E26AD" box="[844,950,937,960]" pageId="20" pageNumber="20" type="holotype">Holotype</typeStatus>
.
</emphasis>
<specimenCode id="DBC59EA96F3E821BFC50FC25FBE826AD" box="[969,1120,937,960]" pageId="20" pageNumber="20">MNHNP 250</specimenCode>
(
<figureCitation id="13582A576F3E821BFBEEFC25FB4726AD" box="[1143,1231,937,960]" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="20" pageNumber="20">Fig. 8C</figureCitation>
), a nearly complete rostrum without nasals.
</materialsCitation>
</paragraph>
<paragraph id="8BDC36D26F3E821BFCD5FC75FBB3211F" blockId="20.[844,1467,1017,1138]" pageId="20" pageNumber="20">
<emphasis id="B917EAC06F3E821BFCD5FC75FC51217D" bold="true" box="[844,985,1017,1040]" pageId="20" pageNumber="20">Occurrence.</emphasis>
Late Miocene (Messinian), pre-evaporitic period of the Tripoli Unit, locality of Les Planteurs,
<collectingCountry id="F37476426F3E821BFCD5FBB6FC14213C" box="[844,924,1082,1105]" name="Algeria" pageId="20" pageNumber="20">Algeria</collectingCountry>
(
<bibRefCitation id="EFF24B236F3E821BFC2DFBB6FB6F213C" author="Landini, W. &amp; Menesini, E." box="[948,1255,1082,1105]" pageId="20" pageNumber="20" pagination="279 - 290" refId="ref27152" refString="Landini, W. &amp; Menesini, E. 1984. Messinian marine fish communities of the Mediterranean Sea. Atti della Societa`Toscana di Scienze Naturali, Serie A, 91, 279 - 290." type="journal article" year="1984">Landini &amp; Menesini 1984</bibRefCitation>
;
<bibRefCitation id="EFF24B236F3E821BFB66FBB6FC09211F" author="Carnevale, G. &amp; Sorbini, C. &amp; Landini, W. &amp; Varola, A." pageId="20" pageNumber="20" pagination="63 - 75" refId="ref24342" refString="Carnevale, G., Sorbini, C., Landini, W. &amp; Varola, A. 2002. Makaira cf. M. nigricans Lacepede, 1802 (Teleostei: Perciformes: Istiophoridae) from the Pietra Leccese, late Miocene, Apulia, southern Italy. Palaeontographia Italica, 88, 63 - 75." type="journal article" year="2002">
Carnevale
<emphasis id="B917EAC06F3E821BFAE2FBB6FA33213C" box="[1403,1467,1082,1105]" italics="true" pageId="20" pageNumber="20">et al.</emphasis>
2002
</bibRefCitation>
;
<bibRefCitation id="EFF24B236F3E821BFC0BFBD7FBA2211F" author="Gaudant, J." box="[914,1066,1115,1138]" pageId="20" pageNumber="20" pagination="141 - 163" refId="ref25722" refString="Gaudant, J. 2008. Pal´eobiodiversit´e et pal´eoenvironnements: l' exemple des gisements de poissons t´el´eost´eens du Messinien pr´e´evaporitique d' Oran et du bassin du Ch´elif (Alg´erie). Geodiversitas, 30, 141 - 163." type="journal article" year="2008">Gaudant 2008</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37965596F3E8219FCD5FB07FD9C27D8" lastPageId="22" lastPageNumber="22" pageId="20" pageNumber="20" type="discussion">
<paragraph id="8BDC36D26F3E821BFCD5FB07FB80208A" blockId="20.[844,1467,1163,1901]" pageId="20" pageNumber="20">
<emphasis id="B917EAC06F3E821BFCD5FB07FC3321CF" bold="true" box="[844,955,1163,1186]" pageId="20" pageNumber="20">Remarks.</emphasis>
The preserved rostrum is
<quantity id="4C9B9B376F3E821BFB46FB07FABC21CF" box="[1247,1332,1163,1186]" metricMagnitude="-1" metricUnit="m" metricValue="2.95" pageId="20" pageNumber="20" unit="mm" value="295.0">295 mm</quantity>
long, rather slender, with a generally elliptical cross-section (wider than tall) (
<figureCitation id="13582A576F3E821BFC58FB40FB9B218E" box="[961,1043,1228,1251]" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="20" pageNumber="20">Fig. 8C</figureCitation>
). In cross-section, two continuous and large internal internal canals are visible (
<tableCitation id="C6E103696F3E821BFABDFB61FAF32069" box="[1316,1403,1261,1284]" captionStart="Table 3" captionStartId="9.[159,218,187,208]" captionTargetPageId="9" 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)." httpUri="http://table.plazi.org/id/DF1C665A6F238206FF06FF37FEEB2438" pageId="20" pageNumber="20" tableUuid="DF1C665A6F238206FF06FF37FEEB2438">Table 3</tableCitation>
;
<figureCitation id="13582A576F3E821BFA08FB61FCE32049" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="20" pageNumber="20">Fig. 8C</figureCitation>
). In lateral view, the rostrum is straight dorsally, but slightly curved upwards ventrally (
<figureCitation id="13582A576F3E821BFB5EFAA2FA922028" box="[1223,1306,1326,1349]" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="20" pageNumber="20">Fig. 8C</figureCitation>
). It is covered partly by conical denticles that are restricted to the ventral and lateral surfaces of the specimen. The denticles located on the lateral sides are the largest, while those located on the ventral surface are smaller (Supplemental material
<figureCitation id="13582A576F3E821BFC32FA5CFC70208A" box="[939,1016,1488,1511]" captionStart="Figure 4" captionStartId="10.[159,226,1174,1195]" captionTargetBox="[159,1469,191,1149]" captionTargetId="figure-227@10.[159,1469,188,1157]" captionTargetPageId="10" captionText="Figure 4. Maximum likelihood constrained topology based on Williams et al. (2018) for istiophorid billfishes. Colours indicate different clades that are correlated with different types of rostra and behaviours based on extant species. The definition for the rostra shape is detailed in the character matrix (Supplemental material Table S2). The skull figures in ventral view show the morphology of the neurocranium, vomer and parasphenoid in different clades. The colours for skull bones are explained in Figure 13. The rostrum and drawings on the button show how each type of rostra correlated with a different feeding behaviours based on observations in extant species. The silhouettes of the three main types of rostra described are compared with drawings of different types of swords for better understanding of the rostral morphology. Green is extremely thin rostra similar to a fencing sword; blue illustrates robust rostrums compared with a gladius hispaniensis or Mainz-type swords, as shown in Bishop (2016, pp. 1821); red shows slender rostrums compared with a Viking sword (see Oakeshott [1960] for examples of this type of sword) and orange indicates the same rostrum type as red. Colours also show the convergence of different rostral, vomer and parasphenoid shapes in our topology. The Tetrapturus clade (orange) includes species with slender rostra similar to the Gracilorosta clade (red) but no information about the feeding behaviour in spearfishes is available." pageId="20" pageNumber="20">Fig. S4</figureCitation>
).
</paragraph>
<paragraph id="8BDC36D26F3E8219FCFEFA7CFD9C27D8" blockId="20.[844,1467,1163,1901]" lastBlockId="22.[159,781,189,693]" lastPageId="22" lastPageNumber="22" pageId="20" pageNumber="20">
The cross-sections of the rostrum change along its length and each section is described here in the same sequence as in
<bibRefCitation id="EFF24B236F3E821BFB99F9BDFB592325" author="Arambourg, C." box="[1024,1233,1585,1608]" pageId="20" pageNumber="20" pagination="1 - 298" refId="ref23620" refString="Arambourg, C. 1927. Les poissons fossiles d' Oran. Mat´eriaux pour la Carte g´eologiques de l' Alg´erie, S´erie 1, Pal´eontologie, 6, 1 - 298." type="journal article" year="1927">Arambourg (1927)</bibRefCitation>
. The first section is represented by the fracture at the proximal end of the rostrum and is equivalent to the 0.5 L distance (
<figureCitation id="13582A576F3E821BFA09F9FEFCE523C7" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="20" pageNumber="20">Fig. 9A</figureCitation>
). At this distance, the rostrum is oval in cross-section but with an angularly obtuse ventral surface. The shape for the second section (
<figureCitation id="13582A576F3E821BFB06F958FB7C2386" box="[1183,1268,1748,1771]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="20" pageNumber="20">Fig. 9B</figureCitation>
) is similar to the first section. The obtuse ventral angle forms a convex surface along the ventral surface of the rostrum that decreases gradually as it advances distally up to approximately
<quantity id="4C9B9B376F3E821BFC6AF8DAFBC02200" box="[1011,1096,1878,1901]" metricMagnitude="-1" metricUnit="m" metricValue="1.39" pageId="20" pageNumber="20" unit="mm" value="139.0">139 mm</quantity>
(if measured from the proximal end) (
<figureCitation id="13582A576F3C8219FF7BFF31FED525B9" box="[226,349,189,212]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9A, B</figureCitation>
). At this distance, the third section is located and is equivalent to that at 0.25 L (
<figureCitation id="13582A576F3C8219FD3CFF51FD752599" box="[677,765,221,244]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9C</figureCitation>
). Here the rostrum is oval and two smaller canals located dorsolateral to the centre of the bone are visible (
<figureCitation id="13582A576F3C8219FD7AFE91FF362439" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9C</figureCitation>
). These small canals are visible for about
<quantity id="4C9B9B376F3C8219FD5DFEB1FC842439" box="[708,780,317,340]" metricMagnitude="-2" metricUnit="m" metricValue="6.2" pageId="22" pageNumber="22" unit="mm" value="62.0">62 mm</quantity>
from this point extending dorsally before they gradually disappear distally. The fourth section is located at
<quantity id="4C9B9B376F3C8219FF06FE11FF7C24D9" box="[159,244,413,436]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="22" pageNumber="22" unit="mm" value="200.0">200 mm</quantity>
(
<figureCitation id="13582A576F3C8219FE93FE11FEE924D9" box="[266,353,413,436]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9D</figureCitation>
). It is oval but with a planar ventral margin and the second pair of internal canals are visible (
<figureCitation id="13582A576F3C8219FF31FE51FF752499" box="[168,253,477,500]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9D</figureCitation>
). From this point (at
<quantity id="4C9B9B376F3C8219FE6AFE51FDC22499" box="[499,586,477,500]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="22" pageNumber="22" unit="mm" value="200.0">200 mm</quantity>
) to the distal tip, the ventral planar surface extends for about
<quantity id="4C9B9B376F3C8219FD1DFE72FD472778" box="[644,719,510,533]" metricMagnitude="-2" metricUnit="m" metricValue="4.6" pageId="22" pageNumber="22" unit="mm" value="46.0">46 mm</quantity>
, until the rostrum acquires an oval shape anew in cross-section. The last section is located at
<quantity id="4C9B9B376F3C8219FDA2FDB2FD182738" box="[571,656,574,597]" metricMagnitude="-1" metricUnit="m" metricValue="2.6" pageId="22" pageNumber="22" unit="mm" value="260.0">260 mm</quantity>
(
<figureCitation id="13582A576F3C8219FD31FDB2FD752738" box="[680,765,574,597]" 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-section approximately at 0.5L distance (prenasal sutures are not shown due to the poor preservation state). H, I, N, †Makaira belgica, MAUL 917/1; H, cross-section at 0.5L and I, 0.25 L distance; IRSNB P117; N, cross-section at 0.25 L distance (prenasal sutures are not shown due to the poor preservation). J, K, †Spathochoira calvertense gen et. comb. nov. (USNM 9344) at J, 0.5 L and K, 0.25 L; L, M, extant I. platypterus at L, 0.5L and M, 0.25 L; O, †Xiphiorhynchus solidus (IRSNB P643), cross-section at the proximal end of the rostrum. Dashed lines show reconstruction of missing sections. The black arrow in I, shows the dorsal placement of the internal canals. In J, K, the arrows show the ventral external canal in cross-section at 0.5 L and 0.25L distance. Scale bars = 1 cm." pageId="22" pageNumber="22">Fig. 9E</figureCitation>
). Here the lateral sides (when observed in cross-section) are compressed and the distal tip acquires a slender and very pointed shape (
<figureCitation id="13582A576F3C8219FEE7FD12FE5E27D8" box="[382,470,670,693]" 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="22" pageNumber="22">Figs 7A</figureCitation>
,
<figureCitation id="13582A576F3C8219FE7CFD12FD8D27D8" box="[485,517,670,693]" captionStart="Figure 8" captionStartId="16.[159,226,1150,1171]" captionTargetBox="[177,1469,197,1132]" captionTargetId="figure-269@16.[159,1469,188,1133]" captionTargetPageId="16" captionText="Figure 8. AD, Istiophoridae holotypes represented by fragmented rostrums in dorsal, right lateral and ventral view; A, †Spathochoira calvertense gen et. comb. nov. (USNM 9344), holotype; B, Makaira belgica (IRSNB P1117), holotype; C, †Prototetrapturus courcelli gen et. comb. nov. (MNHN P250), holotype; D, Makaira purdyi (USNM 481933), holotype. Dashed black lines show the positions where the sections at 0.5L and 0.25L have been studied. White dashed lines indicates the prenasal sutures. Scale bar = 5 cm." pageId="22" pageNumber="22">8C</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>