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<document id="04C7D6987F8777F091498A86EE4C7291" ID-DOI="10.1080/14772019.2018.1446462" ID-ISSN="1478-0941" ID-PMC="PMC6474738" ID-PubMed="31057335" ID-Zenodo-Dep="10883523" 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="1711484535305" checkinUser="felipe" docAuthor="Torres-Silva, Ana. I., Eder, Wolfgang, Hohenegger, Johann &amp; Briguglio, Antonino" docDate="2018" docId="9E6A87F1FFC5282CFC42F90395D28BE5" docLanguage="en" docName="JSystPalaeontol.17.7.557-595.pdf" docOrigin="Journal of Systematic Palaeontology 17 (7)" docSource="http://dx.doi.org/10.1080/14772019.2018.1446462" docStyle="DocumentStyle:602474738E3D100F0F13788B53F61545.2:JSystPalaeontol.2016-.journal_article.open" docStyleId="602474738E3D100F0F13788B53F61545" docStyleName="JSystPalaeontol.2016-.journal_article.open" docStyleVersion="2" docTitle="Heterostegina ocalana Cushman 1921" docType="treatment" docVersion="3" lastPageNumber="587" masterDocId="6253FF89FFDB2833FFF3FFF491548914" masterDocTitle="Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends" masterLastPageNumber="595" masterPageNumber="557" pageNumber="586" updateTime="1711485251774" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="FCE24881A0DDAE62F2F3340D5F5C0BB5">Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends</mods:title>
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<mods:namePart id="B6B88315F4F99953F2B68FF5095F17F8">Torres-Silva, Ana. I.</mods:namePart>
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<mods:namePart id="B2AD4A699D8752A6C99517F14B0C3BCB">Eder, Wolfgang</mods:namePart>
<mods:affiliation id="ACC6A312EEE4837145CA9005A6A9B9AF">Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria;</mods:affiliation>
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<mods:namePart id="B882DBAE1DB7395F6AD61DD3A9142592">Hohenegger, Johann</mods:namePart>
<mods:affiliation id="26F833B329EECA8BA949D980A5ADE19B">Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090, Vienna, Austria;</mods:affiliation>
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<mods:namePart id="D115494BECE8112F8869FA6F86E25D99">Briguglio, Antonino</mods:namePart>
<mods:affiliation id="EE82C024D0C34C4F5785D3689138F6C1">Dipartimento di Scienze della, Terra dellAmbiente e della Vita, Universita ́ degli Studi di Genova, Corso Europa, 26, I- 16132 Genova, Italy</mods:affiliation>
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<subSubSection id="5ED9656CFFC5282DFC42F90394028E1A" box="[945,1366,1783,1807]" pageId="30" pageNumber="586" type="nomenclature">
<paragraph id="167C36E7FFC5282DFC42F90394028E1A" blockId="30.[945,1366,1783,1838]" box="[945,1366,1783,1807]" pageId="30" pageNumber="586">
<heading id="4D34818BFFC5282DFC42F90394028E1A" box="[945,1366,1783,1807]" centered="true" fontSize="10" level="2" pageId="30" pageNumber="586" reason="8">
<taxonomicName id="D1C34D64FFC5282DFC42F90394028E1A" ID-CoL="6LTT8" authority="Cushman, 1921" authorityName="Cushman" authorityYear="1921" box="[945,1366,1783,1807]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="30" pageNumber="586" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC5282DFC42F90395FD8E1B" bold="true" box="[945,1193,1783,1807]" italics="true" pageId="30" pageNumber="586">Heterostegina ocalana</emphasis>
<bibRefCitation id="72524B16FFC5282DFB5CF90394028E1A" author="Cushman, J. A." box="[1199,1366,1783,1806]" pageId="30" pageNumber="586" pagination="125 - 137" refId="ref20551" refString="Cushman, J. A. 1921. American species of Operculina and Heterostegina and their faunal relations. US Geological Survey Professional Paper, 128 - E, 125 - 137." type="book chapter" year="1921">Cushman, 1921</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="5ED9656CFFC5282CFBEFF8E3906C8B47" lastPageId="31" lastPageNumber="587" pageId="30" pageNumber="586" type="description">
<paragraph id="167C36E7FFC5282DFBEFF8E395BF8E3A" blockId="30.[945,1366,1783,1838]" box="[1052,1259,1815,1838]" pageId="30" pageNumber="586">
(
<figureCitation id="8EF82A62FFC5282DFBD6F8E395B08E3A" box="[1061,1252,1815,1838]" captionStart="Figure 18" captionStartId="30.[159,227,1281,1302]" captionTargetBox="[175,1446,185,1247]" captionTargetId="figure-178@30.[173,1453,183,1249]" captionTargetPageId="30" captionText="Figure 18. A, B, DH, Heterostegina ocalana Cushman; A, Loma Viǵıa, CA-216-D1a; B, Norona, NOR-UN 15/14; D, Loma Viǵıa, CA-216-79; E, F, Loma Jabaco; E, LM-52-756; F, LM-52-752; G, H, Norona, NOR-UN 24. C, Heterostegina cubana Cizancourt, Loma candelaria, 98LC-1H-809. I, Heterostegina sp. indet., Loma Candelaria, 98LC-1H-808. A, B, A forms in axial section; CG, I, A forms in equatorial section; H, external view." figureDoi="http://doi.org/10.5281/zenodo.10883565" httpUri="https://zenodo.org/record/10883565/files/figure.png" pageId="30" pageNumber="586">Fig. 18A, B, DH</figureCitation>
)
</paragraph>
<paragraph id="167C36E7FFC5282DFCBEF8A392E58E9A" blockId="30.[845,1466,1879,1934]" pageId="30" pageNumber="586">
1921
<taxonomicName id="D1C34D64FFC5282DFC7EF8A395AC8E7A" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[909,1272,1879,1902]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="30" pageNumber="586" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC5282DFC7EF8A395D28E7A" box="[909,1158,1879,1902]" italics="true" pageId="30" pageNumber="586">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
, 130, pl. 21, figs 1518.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFF49904189E0" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
1941
<taxonomicName id="D1C34D64FFC4282CFF2CFF49931C89C0" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[223,584,189,212]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFF2CFF49908289C0" box="[223,470,189,212]" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
; Cole: 32, pl. 11, figs 36.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFF0990778820" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
1952
<taxonomicName id="D1C34D64FFC4282CFF12FF09931A8800" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[225,590,253,276]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFF12FF09908E8800" box="[225,474,253,276]" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
; Cole: 13, pl. 4, figs 218.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFEC990E88860" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
1957
<taxonomicName id="D1C34D64FFC4282CFF13FEC9931F8840" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[224,587,317,340]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFF13FEC9908C8840" box="[224,472,317,340]" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
; Puri: 136, pl. 6, figs 10, 11, pl. 7, fig. 16.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFE89934188A0" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
1993
<taxonomicName id="D1C34D64FFC4282CFF2EFE8993E38880" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[221,695,381,404]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana" subGenus="Vlerkina">
<emphasis id="24B7EAF5FFC4282CFF2EFE8993138880" box="[221,583,381,404]" italics="true" pageId="31" pageNumber="587">Heterostegina (Vlerkina) ocalana</emphasis>
Cushman
</taxonomicName>
; Robinson &amp; Wright: 335, pl. 31, fig. 4.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFE48925188E7" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
2017
<taxonomicName id="D1C34D64FFC4282CFF1AFE48933C88C7" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[233,616,444,467]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFF1AFE4890BF88C7" box="[233,491,444,467]" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
; Torres-Silva, Hohenegger, Cori ́́c, Briguglio, &amp; Eder: 57, fig. 10BD.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CFE08906C8B47" blockId="31.[159,781,189,595]" pageId="31" pageNumber="587">
2017
<taxonomicName id="D1C34D64FFC4282CFF13FE0893198B07" authority="Cushman" authorityName="Cushman" authorityYear="1921" box="[224,589,508,531]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFF13FE08908D8B07" box="[224,473,508,531]" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
Cushman
</taxonomicName>
; Benedetti, Less, Parente, Pignatti, Cahizac, Torres-Silva, &amp; Buhl: 14, fig. 10AG.
</paragraph>
</subSubSection>
<subSubSection id="5ED9656CFFC4282CFF6CFD9890188BF6" pageId="31" pageNumber="587" type="materials_examined">
<paragraph id="167C36E7FFC4282CFF6CFD9890188BF6" blockId="31.[159,781,620,738]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFF6CFD98905E8B97" bold="true" box="[159,266,620,643]" pageId="31" pageNumber="587">Material.</emphasis>
Twenty-seven megalospheric specimens in equatorial sections, comprising 14 from Loma Viǵıa (CA-216), nine from Loma Jabaco (LM-52) and four from Norona (
<collectionCode id="70D2AE22FFC4282CFF27FD3F90168BF6" box="[212,322,715,738]" pageId="31" pageNumber="587">NOR-UN</collectionCode>
).
</paragraph>
</subSubSection>
<subSubSection id="5ED9656CFFC4282CFF6CFD0F930E8E9A" pageId="31" pageNumber="587" type="description">
<paragraph id="167C36E7FFC4282CFF6CFD0F907E8A06" blockId="31.[159,781,763,1169]" box="[159,298,763,786]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFF6CFD0F907E8A06" bold="true" box="[159,298,763,786]" pageId="31" pageNumber="587">Description.</emphasis>
</paragraph>
<paragraph id="167C36E7FFC4282CFF34FCEF90A38D85" blockId="31.[159,781,763,1169]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFF34FCEF90C88A26" bold="true" box="[199,412,795,818]" pageId="31" pageNumber="587">External features.</emphasis>
The test is involute, becoming evolute in the last whorls, lenticular to flat, biconvex, and thin towards the periphery with oval contour. Tests of the megalospheric form range in diameter from 1.5 to 4.5 mm. The distinct central pile is situated near the embryonic chambers. The septal sutures are slightly curved, and towards the periphery the primary and secondary sutures form a characteristic reticulate network; this ornamentation is absent in the Loma Vigia populations. B forms are very rare and no significant difference was observed in the size of adults between the megalospheric and microspheric forms.
</paragraph>
<paragraph id="167C36E7FFC4282CFF34FB5E91B98E0A" blockId="31.[159,782,1194,1822]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFF34FB5E90F48DD5" bold="true" box="[199,416,1194,1217]" pageId="31" pageNumber="587">Internal features.</emphasis>
Megalospheric generations in equatorial section are characterized by a small and subspherical proloculus 0.0670.19 mm in diameter (mean 0.14 mm), followed by a second reniform chamber and a loosely coiled spiral. The number of post-embryonic undivided chambers ranges from two to four and they do not reappear after the first heterosteginid chamber. Primary septa, with weaker backbend angle, form arched chambers, subdivided into subrectangular chamberlets by complete septula. The first chamberlets along the inner spiral cord are 2 times wider than the others. The number of chamberlets and septula increases through ontogeny. A second megalospheric morphotype was found with proloculus size between 0.05 and 0.06 mm followed by 67 operculinid chambers, confirming the results of
<bibRefCitation id="72524B16FFC4282CFD6CF99C91A68F8B" author="Eder, W. &amp; Hohenegger, J. &amp; Briguglio, A." pageId="31" pageNumber="587" pagination="110 - 117" refId="ref20720" refString="Eder, W., Hohenegger, J. &amp; Briguglio, A. 2017 a. Depthrelated morphoclines of megalospheric tests of Heterostegina depresa d'Orbigny: biostratigraphic and paleobiological implications. Palaios, 32, 110 - 117." type="journal article" year="2017">
Eder
<emphasis id="24B7EAF5FFC4282CFD2BF99C92538F6B" box="[728,775,1640,1663]" italics="true" pageId="31" pageNumber="587">et al</emphasis>
. (2017a)
</bibRefCitation>
on the extant
<taxonomicName id="D1C34D64FFC4282CFE59F97C93F08F8B" authorityName="d'Orbigny" authorityYear="1826" box="[426,676,1672,1695]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="depresa">
<emphasis id="24B7EAF5FFC4282CFE59F97C93F08F8B" box="[426,676,1672,1695]" italics="true" pageId="31" pageNumber="587">Heterostegina depresa</emphasis>
</taxonomicName>
showing strong variability in both characters based on the mixture of two megalospheric generations. Morphological variability in
<taxonomicName id="D1C34D64FFC4282CFEFAF91390D58FEA" authorityName="Cushman" authorityYear="1921" box="[265,385,1767,1790]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFEFAF91390D58FEA" box="[265,385,1767,1790]" italics="true" pageId="31" pageNumber="587">H. ocalana</emphasis>
</taxonomicName>
has already been published by
<bibRefCitation id="72524B16FFC4282CFD28F91391BC8E0A" author="Cole, W. S." pageId="31" pageNumber="587" pagination="748 - 749" refId="ref20169" refString="Cole, W. S. 1953. Larger Foraminifera from the upper Eocene of Costa Rica. Journal of Paleontology, 27, 748 - 749." type="journal article" year="1953">Cole (1953)</bibRefCitation>
.
</paragraph>
<paragraph id="167C36E7FFC4282CFF6CF8C3930E8E9A" blockId="31.[159,781,1847,1934]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFF6CF8C390628E5A" bold="true" box="[159,310,1847,1870]" pageId="31" pageNumber="587">Occurrences.</emphasis>
Late Eocene, NP17/19, Blanco Formation; late Eocene, NP19-20/CP 15 and?early Oligocene, O1/ P18 and NP 21/CP 16, Jabaco Formation.
</paragraph>
</subSubSection>
<subSubSection id="5ED9656CFFC4282CFCBEFF4995D28BE5" pageId="31" pageNumber="587" type="discussion">
<paragraph id="167C36E7FFC4282CFCBEFF4994318B67" blockId="31.[845,1467,189,627]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFCBEFF4992EF89C0" bold="true" box="[845,955,189,212]" pageId="31" pageNumber="587">Remarks.</emphasis>
As the most widely recorded heterostegenid species in the American-Caribbean late Eocene,
<taxonomicName id="D1C34D64FFC4282CFA9BFF2992A38800" authorityName="Cushman" authorityYear="1921" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFA9BFF2992A38800" italics="true" pageId="31" pageNumber="587">Heterostegina ocalana</emphasis>
</taxonomicName>
is distinguished by its small proloculi and the great variability in the number of operculinid chambers within specimens from different localities (Cole 1952;
<bibRefCitation id="72524B16FFC4282CFC67FEA995C88860" author="Torres-Silva, A. I. &amp; Hohenegger, J. &amp; Briguglio, A. &amp; Eder, W." box="[916,1180,349,372]" pageId="31" pageNumber="587" pagination="44 - 60" refId="ref23207" refString="Torres-Silva, A. I., Hohenegger, J., Cori´c´, S., Briguglio, A. &amp; Eder, W. 2017. Biostratigraphy and evolutionary tendencies of Eocene heterostegines in Western and Central Cuba based on morphomteric analyses. Palaios, 32, 44 - 60." type="journal article" year="2017">
Torres-Silva
<emphasis id="24B7EAF5FFC4282CFBD7FEA995038860" box="[1060,1111,349,372]" italics="true" pageId="31" pageNumber="587">et al</emphasis>
. 2017
</bibRefCitation>
). Occurrences of
<taxonomicName id="D1C34D64FFC4282CFA94FEA992288880" authorityName="Cushman" authorityYear="1921" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFA94FEA992288880" italics="true" pageId="31" pageNumber="587">H. ocalana</emphasis>
</taxonomicName>
at Dowling Park (
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) dated by strontium isotope stratigraphy correspond to the latest Priabonian, roughly fitting the E16 planktonic foraminiferal biozone and NP21 calcareous nanofossil zone (
<bibRefCitation id="72524B16FFC4282CFB89FE28940F88E7" author="Benedetti, A. &amp; Less, G. &amp; Parente, M. &amp; Pignatti, J. &amp; Cahizac, B. &amp; Torres-Silva, A. I. &amp; Buhl, D." box="[1146,1371,476,499]" pageId="31" pageNumber="587" pagination="87 - 110" refId="ref19129" refString="Benedetti, A., Less, G., Parente, M., Pignatti, J., Cahizac, B., Torres-Silva, A. I. &amp; Buhl, D. 2017. Heterostegina matteuccii sp. nov. (Foraminiferida: Nummulitidae) from the lower Oligocene of Sicily and Aquitaine: a possible transatlantic inmigrant. Journal of Systematic Palaeontology, 16, 87 - 110." type="journal article" year="2017">
Benedetti
<emphasis id="24B7EAF5FFC4282CFB1AFE28944D88E7" box="[1257,1305,476,499]" italics="true" pageId="31" pageNumber="587">et al</emphasis>
. 2017
</bibRefCitation>
). This is also consistent with the possible early Rupelian age of samples with
<taxonomicName id="D1C34D64FFC4282CFC1DFDE8953F8B27" authorityName="Cushman" authorityYear="1921" box="[1006,1131,540,563]" class="Globothalamea" family="Nummulitidae" genus="Heterostegina" kingdom="Chromista" order="Rotaliida" pageId="31" pageNumber="587" phylum="Foraminifera" rank="species" species="ocalana">
<emphasis id="24B7EAF5FFC4282CFC1DFDE8953F8B27" box="[1006,1131,540,563]" italics="true" pageId="31" pageNumber="587">H. ocalana</emphasis>
</taxonomicName>
from the Norona section, in which this species appears to be significantly more highly evolved than the Priabonian specimens elsewhere.
</paragraph>
<paragraph id="167C36E7FFC4282CFCBEFD6E95D28BE5" blockId="31.[845,1467,666,753]" pageId="31" pageNumber="587">
<emphasis id="24B7EAF5FFC4282CFCBEFD6E94298BA5" bold="true" box="[845,1405,666,689]" pageId="31" pageNumber="587">Stratigraphical and geographical distribution.</emphasis>
Late Eocene (Priabonian);
<collectingCountry id="6ED47677FFC4282CFBB6FD4E95D48BC5" box="[1093,1152,698,721]" name="Cuba" pageId="31" pageNumber="587">Cuba</collectingCountry>
, Florida,
<collectingCountry id="6ED47677FFC4282CFB07FD4E941F8BC5" box="[1268,1355,698,721]" name="Panama" pageId="31" pageNumber="587">Panama</collectingCountry>
,
<collectingCountry id="6ED47677FFC4282CFAADFD4E94E38BC5" box="[1374,1463,698,721]" name="Jamaica" pageId="31" pageNumber="587">Jamaica</collectingCountry>
, and Island of
<collectingCountry id="6ED47677FFC4282CFC2DFD2E95D68BE5" box="[990,1154,730,753]" name="Saint Vincent and the Grenadines" pageId="31" pageNumber="587">the Grenadines</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>