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  <front>
    <journal-meta><journal-id journal-id-type="publisher">JM</journal-id><journal-title-group>
    <journal-title>Journal of Micropalaeontology</journal-title>
    <abbrev-journal-title abbrev-type="publisher">JM</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">J. Micropalaeontol.</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">2041-4978</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/jm-40-145-2021</article-id><title-group><article-title>Upper Eocene planktonic foraminifera from northern Saudi Arabia:
implications for stratigraphic ranges</article-title><alt-title>Upper Eocene planktonic foraminifera from northern Saudi Arabia</alt-title>
      </title-group><?xmltex \runningtitle{Upper Eocene planktonic foraminifera from northern Saudi Arabia}?><?xmltex \runningauthor{B. S. Wade et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Wade</surname><given-names>Bridget S.</given-names></name>
          <email>b.wade@ucl.ac.uk</email>
        <ext-link>https://orcid.org/0000-0002-7245-8614</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Aljahdali</surname><given-names>Mohammed H.</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-8194-8810</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>Mufrreh</surname><given-names>Yahya A.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff4">
          <name><surname>Memesh</surname><given-names>Abdullah M.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>AlSoubhi</surname><given-names>Salih A.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>Zalmout</surname><given-names>Iyad S.</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>Department of Earth Sciences, University College London, Gower Street,
London, WC1E 6BT, UK</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Marine Geology Department, Faculty of Marine Sciences,<?xmltex \hack{\break}?> King Abdulaziz
University, P.O. Box 80200, Jeddah 21589, Saudi Arabia</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Saudi Geological Survey, Department of Sedimentary Geology, Jeddah,
Saudi Arabia</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Saudi Geological Survey, Department of Paleontology, Jeddah, Saudi
Arabia</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Bridget S. Wade (b.wade@ucl.ac.uk)</corresp></author-notes><pub-date><day>28</day><month>September</month><year>2021</year></pub-date>
      
      <volume>40</volume>
      <issue>2</issue>
      <fpage>145</fpage><lpage>161</lpage>
      <history>
        <date date-type="received"><day>7</day><month>February</month><year>2021</year></date>
           <date date-type="rev-recd"><day>20</day><month>August</month><year>2021</year></date>
           <date date-type="accepted"><day>29</day><month>August</month><year>2021</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2021 Bridget S. Wade et al.</copyright-statement>
        <copyright-year>2021</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021.html">This article is available from https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021.html</self-uri><self-uri xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021.pdf">The full text article is available as a PDF file from https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021.pdf</self-uri>
      <abstract><title>Abstract</title>
    <p id="d1e146">The Rashrashiyah Formation of the Sirhan Basin in northern Saudi
Arabia contains diverse assemblages of planktonic foraminifera. We examined
the biostratigraphy, stratigraphic range and preservation of upper Eocene
planktonic foraminifera. Assemblages are well-preserved and diverse, with
40 species and 11 genera. All samples are assigned to the Priabonian
<italic>Globigerinatheka semiinvoluta</italic> Highest Occurrence Zone (E14), consistent with calcareous nannofossil
biostratigraphy indicating Zone CNE17. Well-preserved planktonic
foraminifera assemblages from the lower part of the upper Eocene are rare
worldwide. Our study provides new insights into the stratigraphic ranges of
many species. We find older (Zone E14) stratigraphic occurrences of several
species of <italic>Globoturborotalita</italic> previously thought to have evolved in the latest Eocene (Zone
E15, E16) or Oligocene; these include <italic>G. barbula, G. cancellata, G. gnaucki, G. pseudopraebulloides</italic>, and <italic>G. paracancellata</italic>. Older stratigraphic
occurrences for <italic>Dentoglobigerina taci</italic> and <italic>Subbotina projecta</italic> are also found, and <italic>Globigerinatheka kugleri</italic> occurs at a younger stratigraphic
level than previously proposed. Our revisions to stratigraphic ranges
indicate that the late Eocene had a higher tropical–subtropical diversity of
planktonic foraminifera than hitherto reported.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d1e180">Planktonic foraminifera are calcareous marine zooplankton. Their abundance
and distinctive morphologies have left a long and valuable marine fossil
record, making them ideal for studies in evolution, climate, and
relationships between diversity and climate change. The diversity of
planktonic foraminifera is high during the middle Eocene (Pearson et al.,
2006; Aze et al., 2011; Fraass et al., 2015), but calcareous and siliceous
zooplankton suffered extinction at the end of the Bartonian associated with
the middle–late Eocene transition (MLET) (Wade, 2004; Kamikuri and Wade,
2012; Wade et al., 2012) and again across the Eocene–Oligocene transition
(EOT) (Wade and Pearson, 2008; Moore and Kamikuri, 2012). However, to date
there have been very few sections with well-preserved lower Priabonian (Zone
E14) planktonic foraminifera assemblages that allow both the wall textures
of specimens to be examined and the full diversity to be documented.</p>
      <p id="d1e183">The term <italic>Konservat-Lagerstätte</italic> (Seilacher, 1970) is used to
characterize exceptional preservation in the fossil record. The Paleogene
sediments of coastal Tanzania contain remarkably well-preserved calcareous
microfossils (Pearson et al., 2001, 2007; Wade and Pearson, 2008), and these
were described as a microfossil <italic>Konservat-Lagerstätte</italic> by Bown et al. (2008). The development of a calcareous microfossil
<italic>Konservat-Lagerstätte</italic> appears to be related to clay-rich sediments that
have never been deeply buried (Bown et al., 2008). The clays act as a low-permeability and low-porosity medium, isolating microfossils from chemical
and physical<?pagebreak page146?> processes of diagenetic alteration. Evidence for exceptional
preservation comes from taxonomic, morphological and geochemical data.
Planktonic foraminifera appear translucent or glassy in reflected light, and
their wall texture is smooth in scanning electron microscope (SEM) images.
These assemblages are probably the nearest approximation to the original
biodiversity, providing a unique snapshot of ancient ecosystems. Planktonic
foraminifera preservation is important to reveal details of wall texture,
imperative for classification and thus phylogeny (e.g., Hemleben and Olsson,
2006). As wall texture underpins genus and higher level taxonomy, only with
high-quality preservation of all the morphological characteristics can
comprehensive assessments of the full diversity be achieved. A lack of
recrystallization and infilling allows delicate features to be observed, for
example intact apertural “teeth” in dentoglobigerinids, the ability to
detect spine holes and therefore distinguish spinose lineages (Pearson and
Wade, 2015; Fayolle and Wade, 2021). It is common to observe a decrease in
the preservation state of buried microfossils. In contrast to “glassy”
preservation, recrystallized specimens appear chalky or white in reflected
light. Diagenesis in planktonic foraminifera can involve overgrowth, changes
in the test crystal structure at the micrometer scale, and/or infilling of the
original test, all of which have significant implications for geochemical
and taxonomic studies.</p>
      <p id="d1e195">While the microfossil Lagerstätten preservation from Tanzania has led to
important insights into the Eocene calcareous microfossil preservation (Bown
et al., 2008), paleoecology and diversity (Wade and Pearson, 2008; Dunkley
Jones et al., 2008), and paleoclimates (Pearson et al., 2001, 2007, 2008,
2009; Lear et al., 2008), Tanzania is not unique in offering such
exceptional preservation. Other sites exist, though their records are not
always as continuous (e.g., Alabama, Miller et al., 2008; Ocean Drilling
Program Site 647, Firth et al., 2013; Integrated Ocean Drilling Program
Expedition 342, Norris et al., 2014). Paleogene sites that contain
excellently preserved microfossils provide significant insights into
planktonic foraminiferal taxonomy and evolutionary snapshots into planktonic
foraminiferal history. High-resolution SEM analysis of well-preserved
planktonic foraminifera can reveal primary wall fabrics that have not
previously been observed. Detailed taxonomic studies are critical to
understanding the phylogeny and evolution of planktonic foraminifera through
the late Eocene. The Tanzania cores, however, do not have recovery in the
lower part of the upper Eocene (Priabonian), as such planktonic foraminifera
taxonomic work focused on the middle Eocene (Pearson et al., 2006) and EOT
(Pearson and Wade, 2015). Here we present new biostratigraphic results and
taxonomic insights from well-preserved material from Saudi Arabia.</p>
      <p id="d1e198">The Paleogene Planktonic Foraminifera Working Group described 14 new
taxa as part of the <italic>Atlas of Oligocene Planktonic Foraminifera</italic> (Wade et al., 2018a). These included four new species
described from the late Eocene and early Oligocene: <italic>Globigerina archaeobulloides</italic> Hemleben and Olsson,
<italic>Globoturborotalita paracancellata</italic> Olsson and Hemleben, <italic>Globoturborotalita pseudopraebulloides</italic> Olsson and Hemleben, and <italic>Subbotina projecta</italic> Olsson, Pearson and Wade.
However, the early stratigraphic ranges of these species were not well
constrained, requiring examination of well-preserved upper Eocene sediments
to determine if these species were present. Our new study section with
well-preserved planktonic foraminifera from Saudi Arabia allows us to
document the presence or absence of these newly described species and
provide constraints to the stratigraphic ranges.</p>
      <p id="d1e217">The calcareous nannofossil biostratigraphy from the upper Eocene
Rashrashiyah Formation in northern Saudi Arabia was recently published in
Aljahdali et al. (2020). Here we present a pilot investigation of the
planktonic foraminifera results for the same section. Whilst our sample set
is limited, we present 96 stacked light microscope and SEM images of the
diverse late Eocene assemblages. Our taxonomic investigations utilize the
<italic>Atlas of Oligocene Planktonic Foraminifera</italic> (Wade et al., 2018a), and we find several species recently described in that
volume but not previously recorded outside the Oligocene. Planktonic
foraminifera results are integrated with previous calcareous nannofossil
records, and the implications for planktonic foraminiferal stratigraphic
ranges are discussed.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Materials and methods</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Geological setting</title>
      <p id="d1e238">The Sirhan (also known as the Azraq-Sirhan) Basin is a northwest–southeast-oriented, regional antisynclinal structure, located in the northern
Arabian Peninsula (Fig. 1). It is part of the Syrian Arc System (Guiraud et
al., 2001) where subsidence in the early Paleogene Tethyan seafloor resulted
in deposition of carbonate and mixed sediments (Al-Rawi, 2014). The
Rashrashiyah Formation crops out in the eastern flank of the Sirhan Basin
(31.46<inline-formula><mml:math id="M1" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> N, 37.36<inline-formula><mml:math id="M2" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula> E) and consists of upper Eocene chalks, claystone
and limestone (Meissner et al., 1990; Halawani, 2001; Aljahdali et al.,
2020) (Figs. 2 and 3). We investigated the uppermost 10 m of the
Rashrashiyah Formation. Five samples were obtained, with a resolution of one
sample per meter, and assigned the prefix A–E (Figs. 2 and 3, Table 1). To
provide stratigraphic constraints, an additional sample (Sample F) was taken
close to the upper contact boundary between Rashrashiyah and Sirhan
formations. The sampling is the same as in Aljahdali et al. (2020) for
calcareous nannofossils.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1"><?xmltex \currentcnt{1}?><?xmltex \def\figurename{Figure}?><label>Figure 1</label><caption><p id="d1e261">Map showing location of the studied section and geological
contexts of the Rashrashiyah Formation in the north-western part of Saudi
Arabia. <bold>(a)</bold> Geological map of the Qurayyat area showing the extension of
the Rashrashiyah Formation around the Qurayyat Regional Airport (modified
after Aljahdali et al., 2020, and Wallace et al., 1994). <bold>(b)</bold> Location of
the Rashrashiyah Formation exposures in north-western Saudi Arabia (map
produced using Ocean Drilling Stratigraphic Network <uri>https://www.odsn.de/</uri>, last access: 17 August 2021).</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f01.png"/>

        </fig>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><?xmltex \currentcnt{2}?><?xmltex \def\figurename{Figure}?><label>Figure 2</label><caption><p id="d1e281">Photograph of the sampled levels of the locally exposed
Rashrashiyah Formation section near the Saudi–Jordanian boarders. Letters
A–F represent sampling levels (photo courtesy of the Saudi Geological
Survey).</p></caption>
          <?xmltex \igopts{width=341.433071pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f02.jpg"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F3"><?xmltex \currentcnt{3}?><?xmltex \def\figurename{Figure}?><label>Figure 3</label><caption><p id="d1e293">Lithology of the studied Rashrashiyah Formation section. Letters
A–F represent sample levels.</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f03.png"/>

        </fig>

<?xmltex \floatpos{p}?><table-wrap id="Ch1.T1" orientation="landscape"><?xmltex \currentcnt{1}?><label>Table 1</label><caption><p id="d1e305">Preservation and relative abundances of planktonic foraminifera
species in the studied part of the Rashrashiyah Formation. <inline-formula><mml:math id="M3" display="inline"><mml:msup><mml:mi/><mml:mo>*</mml:mo></mml:msup></mml:math></inline-formula> Berggren and
Pearson (2005).</p></caption><oasis:table frame="topbot"><?xmltex \begin{scaleboxenv}{.61}[.61]?><oasis:tgroup cols="49">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
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     <oasis:colspec colnum="8" colname="col8" align="center"/>
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     <oasis:colspec colnum="10" colname="col10" align="center"/>
     <oasis:colspec colnum="11" colname="col11" align="center"/>
     <oasis:colspec colnum="12" colname="col12" align="center"/>
     <oasis:colspec colnum="13" colname="col13" align="center"/>
     <oasis:colspec colnum="14" colname="col14" align="center"/>
     <oasis:colspec colnum="15" colname="col15" align="center"/>
     <oasis:colspec colnum="16" colname="col16" align="center"/>
     <oasis:colspec colnum="17" colname="col17" align="center"/>
     <oasis:colspec colnum="18" colname="col18" align="center"/>
     <oasis:colspec colnum="19" colname="col19" align="center"/>
     <oasis:colspec colnum="20" colname="col20" align="center"/>
     <oasis:colspec colnum="21" colname="col21" align="center"/>
     <oasis:colspec colnum="22" colname="col22" align="center"/>
     <oasis:colspec colnum="23" colname="col23" align="center"/>
     <oasis:colspec colnum="24" colname="col24" align="center"/>
     <oasis:colspec colnum="25" colname="col25" align="center"/>
     <oasis:colspec colnum="26" colname="col26" align="center"/>
     <oasis:colspec colnum="27" colname="col27" align="center"/>
     <oasis:colspec colnum="28" colname="col28" align="center"/>
     <oasis:colspec colnum="29" colname="col29" align="center"/>
     <oasis:colspec colnum="30" colname="col30" align="center"/>
     <oasis:colspec colnum="31" colname="col31" align="center"/>
     <oasis:colspec colnum="32" colname="col32" align="center"/>
     <oasis:colspec colnum="33" colname="col33" align="center"/>
     <oasis:colspec colnum="34" colname="col34" align="center"/>
     <oasis:colspec colnum="35" colname="col35" align="center"/>
     <oasis:colspec colnum="36" colname="col36" align="center"/>
     <oasis:colspec colnum="37" colname="col37" align="center"/>
     <oasis:colspec colnum="38" colname="col38" align="center"/>
     <oasis:colspec colnum="39" colname="col39" align="center"/>
     <oasis:colspec colnum="40" colname="col40" align="center"/>
     <oasis:colspec colnum="41" colname="col41" align="center"/>
     <oasis:colspec colnum="42" colname="col42" align="center"/>
     <oasis:colspec colnum="43" colname="col43" align="center"/>
     <oasis:colspec colnum="44" colname="col44" align="center"/>
     <oasis:colspec colnum="45" colname="col45" align="center"/>
     <oasis:colspec colnum="46" colname="col46" align="center"/>
     <oasis:colspec colnum="47" colname="col47" align="center"/>
     <oasis:colspec colnum="48" colname="col48" align="center"/>
     <oasis:colspec colnum="49" colname="col49" align="center"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sample</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col2">Height (m)</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col3">Zone (B&amp;P, 2005<inline-formula><mml:math id="M4" display="inline"><mml:msup><mml:mi/><mml:mo>*</mml:mo></mml:msup></mml:math></inline-formula>)</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col4">Zone (Berggren et al., 1995)</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col5">Preservation</oasis:entry>
         <oasis:entry colname="col6">Notes</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col7">Species richness</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col8"><italic>Acarinina collactea</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col9"><italic>Acarinina echinata</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col10"><italic>Acarinina medizzai</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col11"><italic>Acarinina rohri</italic> (dwarfed)</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col12"><italic>Dentoglobigerina eotripartita</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col13"><italic>Dentoglobigerina galavisi</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col14"><italic>Dentoglobigerina pseudovenezuelana</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col15"><italic>Dentoglobigerina taci</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col16"><italic>Dentoglobigerina tripartita</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col17"><italic>Dentoglobigerina </italic>sp. 1</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col18"><italic>Dentoglobigerina </italic>sp. 2</oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col19"><italic>Globigerina officinalis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col20"><italic>Globigerinatheka barri</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col21"><italic>Globigerinatheka index</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col22"><italic>Globigerinatheka korotkovi</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col23"><italic>Globigerinatheka kugleri</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col24"><italic>Globigerinatheka mexicana</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col25"><italic>Globigerinatheka semiinvoluta</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col26"><italic>Globigerinatheka tropicalis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col27"><italic>Globorotaloides quadrocameratus</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col28"><italic>Globoturborotalita barbula</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col29"><italic>Globoturborotalita cancellata</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col30"><italic>Globoturborotalita gnaucki</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col31"><italic>Globoturborotalita </italic>cf.<italic> G. labiacrassata</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col32"><italic>Globoturborotalita ouachitaensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col33"><italic>Globoturborotalita paracancellata</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col34"><italic>Globoturborotalita pseudopraebulloides</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col35"><italic>Hantkenina alabamensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col36"><italic>Hantkenina primitiva</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col37"><italic>Pseudohastigerina micra</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col38"><italic>Pseudohastigerina naguewichiensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col39"><italic>Subbotina corpulenta</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col40"><italic>Subbotina linaperta</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col41"><italic>Subbotina projecta</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col42"><italic>Subbotina utilisindex</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col43"><italic>Subbotina yeguaensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col44"><italic>Turborotalia ampliapertura</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col45"><italic>Turborotalia cerroazuelensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col46"><italic>Turborotalia cunialensis</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col47"><italic>Turborotalia increbescens</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col48"><italic>Turborotalia pomeroli</italic></oasis:entry>
         <?xmltex \rotentry?><oasis:entry colname="col49"><italic>Turborotalita quinqueloba</italic></oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sarhan F</oasis:entry>
         <oasis:entry colname="col2">9</oasis:entry>
         <oasis:entry colname="col3">–</oasis:entry>
         <oasis:entry colname="col4">–</oasis:entry>
         <oasis:entry colname="col5">VP</oasis:entry>
         <oasis:entry colname="col6">Barren</oasis:entry>
         <oasis:entry colname="col7">0</oasis:entry>
         <oasis:entry colname="col8"/>
         <oasis:entry colname="col9"/>
         <oasis:entry colname="col10"/>
         <oasis:entry colname="col11"/>
         <oasis:entry colname="col12"/>
         <oasis:entry colname="col13"/>
         <oasis:entry colname="col14"/>
         <oasis:entry colname="col15"/>
         <oasis:entry colname="col16"/>
         <oasis:entry colname="col17"/>
         <oasis:entry colname="col18"/>
         <oasis:entry colname="col19"/>
         <oasis:entry colname="col20"/>
         <oasis:entry colname="col21"/>
         <oasis:entry colname="col22"/>
         <oasis:entry colname="col23"/>
         <oasis:entry colname="col24"/>
         <oasis:entry colname="col25"/>
         <oasis:entry colname="col26"/>
         <oasis:entry colname="col27"/>
         <oasis:entry colname="col28"/>
         <oasis:entry colname="col29"/>
         <oasis:entry colname="col30"/>
         <oasis:entry colname="col31"/>
         <oasis:entry colname="col32"/>
         <oasis:entry colname="col33"/>
         <oasis:entry colname="col34"/>
         <oasis:entry colname="col35"/>
         <oasis:entry colname="col36"/>
         <oasis:entry colname="col37"/>
         <oasis:entry colname="col38"/>
         <oasis:entry colname="col39"/>
         <oasis:entry colname="col40"/>
         <oasis:entry colname="col41"/>
         <oasis:entry colname="col42"/>
         <oasis:entry colname="col43"/>
         <oasis:entry colname="col44"/>
         <oasis:entry colname="col45"/>
         <oasis:entry colname="col46"/>
         <oasis:entry colname="col47"/>
         <oasis:entry colname="col48"/>
         <oasis:entry colname="col49"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sarhan E</oasis:entry>
         <oasis:entry colname="col2">4</oasis:entry>
         <oasis:entry colname="col3">E14</oasis:entry>
         <oasis:entry colname="col4">P15</oasis:entry>
         <oasis:entry colname="col5">M</oasis:entry>
         <oasis:entry colname="col6">Specimens are infilled, encrusted, with some<?xmltex \hack{\hfill\break}?>fragmentation</oasis:entry>
         <oasis:entry colname="col7">20</oasis:entry>
         <oasis:entry colname="col8">A</oasis:entry>
         <oasis:entry colname="col9">A</oasis:entry>
         <oasis:entry colname="col10">A</oasis:entry>
         <oasis:entry colname="col11"/>
         <oasis:entry colname="col12">F</oasis:entry>
         <oasis:entry colname="col13">R</oasis:entry>
         <oasis:entry colname="col14">R</oasis:entry>
         <oasis:entry colname="col15">R</oasis:entry>
         <oasis:entry colname="col16"/>
         <oasis:entry colname="col17"/>
         <oasis:entry colname="col18"/>
         <oasis:entry colname="col19"/>
         <oasis:entry colname="col20"/>
         <oasis:entry colname="col21">F</oasis:entry>
         <oasis:entry colname="col22"/>
         <oasis:entry colname="col23"/>
         <oasis:entry colname="col24"/>
         <oasis:entry colname="col25">R</oasis:entry>
         <oasis:entry colname="col26">R</oasis:entry>
         <oasis:entry colname="col27"/>
         <oasis:entry colname="col28"/>
         <oasis:entry colname="col29">R</oasis:entry>
         <oasis:entry colname="col30"/>
         <oasis:entry colname="col31"/>
         <oasis:entry colname="col32">C</oasis:entry>
         <oasis:entry colname="col33"/>
         <oasis:entry colname="col34"/>
         <oasis:entry colname="col35"/>
         <oasis:entry colname="col36">F</oasis:entry>
         <oasis:entry colname="col37">C</oasis:entry>
         <oasis:entry colname="col38"/>
         <oasis:entry colname="col39"/>
         <oasis:entry colname="col40">R</oasis:entry>
         <oasis:entry colname="col41"/>
         <oasis:entry colname="col42">A</oasis:entry>
         <oasis:entry colname="col43"/>
         <oasis:entry colname="col44">R</oasis:entry>
         <oasis:entry colname="col45">F</oasis:entry>
         <oasis:entry colname="col46">R</oasis:entry>
         <oasis:entry colname="col47"/>
         <oasis:entry colname="col48">F</oasis:entry>
         <oasis:entry colname="col49"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sarhan D</oasis:entry>
         <oasis:entry colname="col2">3</oasis:entry>
         <oasis:entry colname="col3">E14</oasis:entry>
         <oasis:entry colname="col4">P15</oasis:entry>
         <oasis:entry colname="col5">G</oasis:entry>
         <oasis:entry colname="col6"/>
         <oasis:entry colname="col7">23</oasis:entry>
         <oasis:entry colname="col8">A</oasis:entry>
         <oasis:entry colname="col9">A</oasis:entry>
         <oasis:entry colname="col10">A</oasis:entry>
         <oasis:entry colname="col11"/>
         <oasis:entry colname="col12"/>
         <oasis:entry colname="col13">R</oasis:entry>
         <oasis:entry colname="col14">R</oasis:entry>
         <oasis:entry colname="col15">F</oasis:entry>
         <oasis:entry colname="col16"/>
         <oasis:entry colname="col17"/>
         <oasis:entry colname="col18"/>
         <oasis:entry colname="col19"/>
         <oasis:entry colname="col20"/>
         <oasis:entry colname="col21">C</oasis:entry>
         <oasis:entry colname="col22"/>
         <oasis:entry colname="col23"/>
         <oasis:entry colname="col24">R</oasis:entry>
         <oasis:entry colname="col25">F</oasis:entry>
         <oasis:entry colname="col26">F</oasis:entry>
         <oasis:entry colname="col27"/>
         <oasis:entry colname="col28"/>
         <oasis:entry colname="col29">R</oasis:entry>
         <oasis:entry colname="col30">A</oasis:entry>
         <oasis:entry colname="col31">F</oasis:entry>
         <oasis:entry colname="col32">C</oasis:entry>
         <oasis:entry colname="col33">F</oasis:entry>
         <oasis:entry colname="col34">C</oasis:entry>
         <oasis:entry colname="col35">R</oasis:entry>
         <oasis:entry colname="col36">F</oasis:entry>
         <oasis:entry colname="col37">C</oasis:entry>
         <oasis:entry colname="col38"/>
         <oasis:entry colname="col39"/>
         <oasis:entry colname="col40"/>
         <oasis:entry colname="col41"/>
         <oasis:entry colname="col42">A</oasis:entry>
         <oasis:entry colname="col43"/>
         <oasis:entry colname="col44">R</oasis:entry>
         <oasis:entry colname="col45"/>
         <oasis:entry colname="col46"/>
         <oasis:entry colname="col47">C</oasis:entry>
         <oasis:entry colname="col48"/>
         <oasis:entry colname="col49">R</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sarhan C</oasis:entry>
         <oasis:entry colname="col2">2</oasis:entry>
         <oasis:entry colname="col3">E14</oasis:entry>
         <oasis:entry colname="col4">P15</oasis:entry>
         <oasis:entry colname="col5">M</oasis:entry>
         <oasis:entry colname="col6">Specimens are infilled</oasis:entry>
         <oasis:entry colname="col7">23</oasis:entry>
         <oasis:entry colname="col8">R</oasis:entry>
         <oasis:entry colname="col9">R</oasis:entry>
         <oasis:entry colname="col10">A</oasis:entry>
         <oasis:entry colname="col11"/>
         <oasis:entry colname="col12">C</oasis:entry>
         <oasis:entry colname="col13"/>
         <oasis:entry colname="col14">R</oasis:entry>
         <oasis:entry colname="col15"/>
         <oasis:entry colname="col16">R</oasis:entry>
         <oasis:entry colname="col17"/>
         <oasis:entry colname="col18"/>
         <oasis:entry colname="col19"/>
         <oasis:entry colname="col20">F</oasis:entry>
         <oasis:entry colname="col21">C</oasis:entry>
         <oasis:entry colname="col22"/>
         <oasis:entry colname="col23"/>
         <oasis:entry colname="col24"/>
         <oasis:entry colname="col25">C</oasis:entry>
         <oasis:entry colname="col26">A</oasis:entry>
         <oasis:entry colname="col27"/>
         <oasis:entry colname="col28"/>
         <oasis:entry colname="col29">R</oasis:entry>
         <oasis:entry colname="col30">A</oasis:entry>
         <oasis:entry colname="col31">F</oasis:entry>
         <oasis:entry colname="col32">C</oasis:entry>
         <oasis:entry colname="col33">F</oasis:entry>
         <oasis:entry colname="col34">C</oasis:entry>
         <oasis:entry colname="col35"/>
         <oasis:entry colname="col36">F</oasis:entry>
         <oasis:entry colname="col37">A</oasis:entry>
         <oasis:entry colname="col38">R</oasis:entry>
         <oasis:entry colname="col39"/>
         <oasis:entry colname="col40"/>
         <oasis:entry colname="col41"/>
         <oasis:entry colname="col42">A</oasis:entry>
         <oasis:entry colname="col43">R</oasis:entry>
         <oasis:entry colname="col44">R</oasis:entry>
         <oasis:entry colname="col45">C</oasis:entry>
         <oasis:entry colname="col46"/>
         <oasis:entry colname="col47"/>
         <oasis:entry colname="col48"/>
         <oasis:entry colname="col49"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Sarhan B</oasis:entry>
         <oasis:entry colname="col2">1</oasis:entry>
         <oasis:entry colname="col3">E14</oasis:entry>
         <oasis:entry colname="col4">P14</oasis:entry>
         <oasis:entry colname="col5">G</oasis:entry>
         <oasis:entry colname="col6">Some fragmentation</oasis:entry>
         <oasis:entry colname="col7">20</oasis:entry>
         <oasis:entry colname="col8">R</oasis:entry>
         <oasis:entry colname="col9">R</oasis:entry>
         <oasis:entry colname="col10">A</oasis:entry>
         <oasis:entry colname="col11"/>
         <oasis:entry colname="col12">C</oasis:entry>
         <oasis:entry colname="col13"/>
         <oasis:entry colname="col14"/>
         <oasis:entry colname="col15"/>
         <oasis:entry colname="col16"/>
         <oasis:entry colname="col17">R</oasis:entry>
         <oasis:entry colname="col18"/>
         <oasis:entry colname="col19"/>
         <oasis:entry colname="col20">F</oasis:entry>
         <oasis:entry colname="col21"/>
         <oasis:entry colname="col22"/>
         <oasis:entry colname="col23">R</oasis:entry>
         <oasis:entry colname="col24"/>
         <oasis:entry colname="col25"/>
         <oasis:entry colname="col26">A</oasis:entry>
         <oasis:entry colname="col27"/>
         <oasis:entry colname="col28">R</oasis:entry>
         <oasis:entry colname="col29"/>
         <oasis:entry colname="col30">A</oasis:entry>
         <oasis:entry colname="col31">F</oasis:entry>
         <oasis:entry colname="col32">C</oasis:entry>
         <oasis:entry colname="col33">F</oasis:entry>
         <oasis:entry colname="col34">A</oasis:entry>
         <oasis:entry colname="col35"/>
         <oasis:entry colname="col36"/>
         <oasis:entry colname="col37">F</oasis:entry>
         <oasis:entry colname="col38"/>
         <oasis:entry colname="col39">F</oasis:entry>
         <oasis:entry colname="col40"/>
         <oasis:entry colname="col41">R</oasis:entry>
         <oasis:entry colname="col42">C</oasis:entry>
         <oasis:entry colname="col43"/>
         <oasis:entry colname="col44"/>
         <oasis:entry colname="col45">A</oasis:entry>
         <oasis:entry colname="col46"/>
         <oasis:entry colname="col47"/>
         <oasis:entry colname="col48">A</oasis:entry>
         <oasis:entry colname="col49"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Sarhan A</oasis:entry>
         <oasis:entry colname="col2">0</oasis:entry>
         <oasis:entry colname="col3">E14</oasis:entry>
         <oasis:entry colname="col4">P14</oasis:entry>
         <oasis:entry colname="col5">VG</oasis:entry>
         <oasis:entry colname="col6">Spines preserved</oasis:entry>
         <oasis:entry colname="col7">27</oasis:entry>
         <oasis:entry colname="col8">F</oasis:entry>
         <oasis:entry colname="col9">F</oasis:entry>
         <oasis:entry colname="col10">F</oasis:entry>
         <oasis:entry colname="col11">R</oasis:entry>
         <oasis:entry colname="col12">F</oasis:entry>
         <oasis:entry colname="col13">R</oasis:entry>
         <oasis:entry colname="col14"/>
         <oasis:entry colname="col15">R</oasis:entry>
         <oasis:entry colname="col16"/>
         <oasis:entry colname="col17"/>
         <oasis:entry colname="col18">R</oasis:entry>
         <oasis:entry colname="col19">F</oasis:entry>
         <oasis:entry colname="col20">C</oasis:entry>
         <oasis:entry colname="col21">F</oasis:entry>
         <oasis:entry colname="col22">R</oasis:entry>
         <oasis:entry colname="col23">R</oasis:entry>
         <oasis:entry colname="col24"/>
         <oasis:entry colname="col25"/>
         <oasis:entry colname="col26">F</oasis:entry>
         <oasis:entry colname="col27">R</oasis:entry>
         <oasis:entry colname="col28">R</oasis:entry>
         <oasis:entry colname="col29">R</oasis:entry>
         <oasis:entry colname="col30"/>
         <oasis:entry colname="col31">F</oasis:entry>
         <oasis:entry colname="col32">C</oasis:entry>
         <oasis:entry colname="col33"/>
         <oasis:entry colname="col34">C</oasis:entry>
         <oasis:entry colname="col35"/>
         <oasis:entry colname="col36"/>
         <oasis:entry colname="col37">F</oasis:entry>
         <oasis:entry colname="col38">R</oasis:entry>
         <oasis:entry colname="col39"/>
         <oasis:entry colname="col40">R</oasis:entry>
         <oasis:entry colname="col41"/>
         <oasis:entry colname="col42">A</oasis:entry>
         <oasis:entry colname="col43"/>
         <oasis:entry colname="col44">R</oasis:entry>
         <oasis:entry colname="col45">F</oasis:entry>
         <oasis:entry colname="col46"/>
         <oasis:entry colname="col47"/>
         <oasis:entry colname="col48">A</oasis:entry>
         <oasis:entry colname="col49"/>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup><?xmltex \end{scaleboxenv}?></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>Sample preparation</title>
      <p id="d1e1519">For planktonic foraminiferal analysis, samples were washed over a 63 <inline-formula><mml:math id="M5" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m
sieve and oven-dried at <inline-formula><mml:math id="M6" display="inline"><mml:mrow><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">40</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M7" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>C. The dried sample was separated
into <inline-formula><mml:math id="M8" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 425, 355–425, 250–355, 150–250 and 63–150 <inline-formula><mml:math id="M9" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m size
fractions, and each size fraction was<?pagebreak page147?> examined under the light microscope.
Following the taxonomy of Pearson et al. (2006), Pearson and Wade (2015), and
Wade et al. (2018a), the abundance of each species was semi-qualitatively
assessed as follows: A – abundant; C – common; F – few; R – rare (Table 1).
Preservation was assessed as VG (very good), where specimens are glassy
under the light microscope with no infilling; G (good), where specimens
are semi-translucent and with no infilling; M (moderate), where specimens
are recrystallized and the test walls are opaque, and VP (very poor),
where specimens are fragmented, opaque and infilled.</p>
      <p id="d1e1564">Selected specimens were picked for <inline-formula><mml:math id="M10" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula>-stacked light microscope and SEM
imaging. A <inline-formula><mml:math id="M11" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula>-stacking light microscope was used to take images in three
views (umbilical, edge and spiral view). For SEM imaging specimens were
placed on stubs<?pagebreak page148?> using double-sided tape. Each stub was coated in an argon
and gold atmosphere using a sputter coater. The illustrated specimens (Figs. 4–9) are deposited in the Natural History Museum, London, UK (NHMUK PM PF
75192–75251).</p>

      <?xmltex \floatpos{p}?><fig id="Ch1.F4" specific-use="star"><?xmltex \currentcnt{4}?><?xmltex \def\figurename{Figure}?><label>Figure 4</label><caption><p id="d1e1583">Light microscope and SEM images of <italic>Acarinina</italic> and <italic>Pseudohastigerina</italic> from the Rashrashiyah
Formation: <bold>(1)</bold> NHMUK PM PF 75192 <italic>Acarinina echinata</italic>, Sample C; <bold>(2a, 2b, 2c)</bold> NHMUK PM PF 75193
<italic>Acarinina medizzai</italic>, Sample A; <bold>(3a, 3b)</bold> NHMUK PM PF 75194 <italic>Acarinina medizzai</italic>, Sample B; <bold>(4a, 4b)</bold> NHMUK PM PF
75195 <italic>Acarinina medizzai</italic>, Sample B; <bold>(5)</bold> NHMUK PM PF 75196 <italic>Acarinina medizzai</italic>, Sample D; <bold>(6)</bold> NHMUK PM PF 75197
<italic>Acarinina medizzai</italic>, Sample A; <bold>(7)</bold> NHMUK PM PF 75231 <italic>Pseudohastigerina naguewichiensis</italic>, Sample C; <bold>(8)</bold> NHMUK PM PF 75232
<italic>Pseudohastigerina naguewichiensis</italic>, Sample A; <bold>(9)</bold> NHMUK PM PF 75233 <italic>Pseudohastigerina micra</italic>, Sample B; <bold>(10)</bold> NHMUK PM PF 75209
<italic>Pseudohastigerina micra</italic>, Sample A; <bold>(11)</bold> NHMUK PM PF 75210 <italic>Pseudohastigerina micra</italic>, Sample A; <bold>(12)</bold> NHMUK PM PF 75211
<italic>Pseudohastigerina micra</italic>, Sample A. Scale bars: <bold>(1)</bold>–<bold>(4a)</bold>, <bold>(5)</bold>–<bold>(12)</bold> <inline-formula><mml:math id="M12" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M13" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m; <bold>(4b)</bold> <inline-formula><mml:math id="M14" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 10 <inline-formula><mml:math id="M15" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f04.jpg"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F5" specific-use="star"><?xmltex \currentcnt{5}?><?xmltex \def\figurename{Figure}?><label>Figure 5</label><caption><p id="d1e1723">Light microscope and SEM images of <italic>Dentoglobigerina</italic> and <italic>Subbotina</italic> from the Rashrashiyah
Formation: <bold>(1)</bold> NHMUK PM PF 75204 <italic>Dentoglobigerina eotripartita</italic>, Sample B; <bold>(2a, 2b)</bold> NHMUK PM PF 75205
<italic>Dentoglobigerina galavisi</italic>, Sample D; <bold>(3)</bold> NHMUK PM PF 75206 <italic>Dentoglobigerina taci</italic>, Sample D; <bold>(4)</bold> NHMUK PM PF 75234
<italic>Dentoglobigerina taci</italic>, Sample A; <bold>(5a, 5b)</bold> NHMUK PM PF 75235 <italic>Dentoglobigerina tripartita</italic>, Sample C; <bold>(6)</bold> NHMUK PM PF 75208
<italic>Dentoglobigerina </italic>sp. 1, Sample B; <bold>(7a, 7b, 7c)</bold> NHMUK PM PF 75207 <italic>Dentoglobigerina </italic>sp. 2, Sample A; <bold>(8a, 8b)</bold> NHMUK PM PF 75212 <italic>Subbotina projecta</italic>, Sample B; <bold>(9a, 9b)</bold> NHMUK PM PF 75213 <italic>Subbotina utilisindex</italic>, Sample B; <bold>(10)</bold> NHMUK PM PF 75214 <italic>Subbotina linaperta</italic>, Sample A. Scale bars: <bold>(1)</bold>–<bold>(9a)</bold>, <bold>(10)</bold> <inline-formula><mml:math id="M16" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M17" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m; <bold>(9b)</bold> <inline-formula><mml:math id="M18" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 10 <inline-formula><mml:math id="M19" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f05.png"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F6" specific-use="star"><?xmltex \currentcnt{6}?><?xmltex \def\figurename{Figure}?><label>Figure 6</label><caption><p id="d1e1847">Light microscope and SEM images of <italic>Subbotina, Hantkenina</italic> and <italic>Globigerinatheka</italic> from the Rashrashiyah
Formation: <bold>(1)</bold> NHMUK PM PF 75236 <italic>Subbotina corpulenta</italic>, Sample B; <bold>(2)</bold> NHMUK PM PF 75199
<italic>Hantkenina primitiva</italic>, Sample C; <bold>(3a, 3b)</bold> NHMUK PM PF 75237 <italic>Globigerinatheka barri</italic>, Sample A; <bold>(4a, 4b)</bold> NHMUK PM PF
75238 <italic>Globigerinatheka barri</italic>, Sample C; <bold>(5a, 5b)</bold> NHMUK PM PF 75239 <italic>Globigerinatheka index</italic>, Sample A; <bold>(6)</bold> NHMUK PM PF
75240 <italic>Globigerinatheka index</italic>, Sample D; <bold>(7a, 7b)</bold> NHMUK PM PF 75241 <italic>Globigerinatheka kugleri,</italic> Sample B; <bold>(8a, 8b)</bold> NHMUK PM PF
75242 <italic>Globigerinatheka kugleri,</italic> Sample A; <bold>(9)</bold> NHMUK PM PF 75243 <italic>Globigerinatheka mexicana</italic>, Sample D; <bold>(10a, 10b)</bold> NHMUK PM PF
75200 <italic>Globigerinatheka mexicana</italic>, Sample D. Scale bars: <bold>(1)</bold>–<bold>(10b)</bold> <inline-formula><mml:math id="M20" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M21" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f06.jpg"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F7" specific-use="star"><?xmltex \currentcnt{7}?><?xmltex \def\figurename{Figure}?><label>Figure 7</label><caption><p id="d1e1949">Light microscope and SEM images of <italic>Globigerinatheka, Globorotaloides </italic>and <italic>Turborotalia </italic>from the Rashrashiyah
Formation: <bold>(1)</bold> NHMUK PM PF 75201 <italic>Globigerinatheka tropicalis</italic>, Sample B; <bold>(2)</bold> NHMUK PM PF 75202
<italic>Globigerinatheka tropicalis</italic>, Sample A; <bold>(3)</bold> NHMUK PM PF 75203 <italic>Globigerinatheka tropicalis</italic>, Sample A; <bold>(4a, 4b)</bold> NHMUK PM PF 75244
<italic>Globigerinatheka semiinvoluta</italic>, Sample C; <bold>(5a, 5b, 5c)</bold> NHMUK PM PF 75245 <italic>Globigerinatheka semiinvoluta</italic>, Sample C; <bold>(6)</bold> NHMUK PM PF 75198
<italic>Globorotaloides quadrocameratus</italic>, Sample A; <bold>(7a, 7b)</bold> NHMUK PM PF 75227 <italic>Turborotalia ampliapertura</italic>, Sample A; <bold>(8)</bold> NHMUK PM PF 75228
<italic>Turborotalia cerroazulensis</italic>, Sample B; <bold>(9a, 9b)</bold> NHMUK PM PF 75229 <italic>Turborotalia cerroazulensis</italic>, Sample A; <bold>(10a, 10b)</bold> NHMUK PM PF
75230 <italic>Turborotalia cerroazulensis</italic>, Sample B. Scale bars: <bold>(1)</bold>–<bold>(10b)</bold> <inline-formula><mml:math id="M22" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M23" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f07.png"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F8" specific-use="star"><?xmltex \currentcnt{8}?><?xmltex \def\figurename{Figure}?><label>Figure 8</label><caption><p id="d1e2051">Light microscope and SEM images of <italic>Globoturborotalita</italic> and <italic>Turborotalita </italic>from the Rashrashiyah
Formation: <bold>(1a, 1b)</bold> NHMUK PM PF 75248 <italic>Globoturborotalita barbula</italic>, Sample A; <bold>(2)</bold> NHMUK PM PF 75249
<italic>Globoturborotalita barbula</italic>, Sample B; <bold>(3)</bold> NHMUK PM PF 75250 <italic>Globoturborotalita cancellata</italic>, Sample A; <bold>(4a, 4b)</bold> NHMUK PM PF 75223
<italic>Globoturborotalita cancellata</italic>, Sample E; <bold>(5a, 5b)</bold> NHMUK PM PF 75224 <italic>Globoturborotalita cancellata</italic>, Sample D; <bold>(6a, 6b)</bold> NHMUK PM PF
75251 <italic>Globoturborotalita gnaucki</italic>, Sample B; <bold>(7a, 7b)</bold> NHMUK PM PF 75225 <italic>Globoturborotalita ouachitaensis</italic>, Sample C; <bold>(8a, 8b, 8c)</bold> NHMUK
PM PF 75226 <italic>Globoturborotalita ouachitaensis</italic>, Sample C; <bold>(9)</bold> NHMUK PM PF 75222 <italic>Turborotalita quinqueloba</italic>, Sample D. Scale bars: <bold>(1)</bold>–<bold>(9)</bold> <inline-formula><mml:math id="M24" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M25" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f08.jpg"/>

        </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F9" specific-use="star"><?xmltex \currentcnt{9}?><?xmltex \def\figurename{Figure}?><label>Figure 9</label><caption><p id="d1e2148">Light microscope and SEM images of <italic>Globoturborotalita</italic> from the Rashrashiyah
Formation: <bold>(1a, 1b)</bold> NHMUK PM PF 75215 <italic>Globoturborotalita </italic>cf. <italic>G. labiacrassata</italic>, Sample D; <bold>(2)</bold> NHMUK PM PF 75216
<italic>Globoturborotalita</italic> cf. <italic>G. labiacrassata</italic>, Sample B; <bold>(3)</bold> NHMUK PM PF 75217 <italic>Globoturborotalita paracancellata</italic>, Sample D; <bold>(4)</bold> NHMUK PM PF 75246
<italic>Globoturborotalita paracancellata</italic>, Sample B; <bold>(5a, 5b, 5c)</bold> NHMUK PM PF 75219 <italic>Globoturborotalita pseudopraebulloides</italic>, Sample B; <bold>(6a, 6b, 6c)</bold> NHMUK PM
PF 75218 <italic>Globoturborotalita pseudopraebulloides</italic>, Sample B; <bold>(7)</bold> NHMUK PM PF 75220 <italic>Globoturborotalita pseudopraebulloides</italic>, Sample A; <bold>(8a, 8b, 8c)</bold> NHMUK PM
PF 75221 <italic>Globoturborotalita pseudopraebulloides</italic>, Sample A; <bold>(9)</bold> NHMUK PM PF 75247 <italic>Globoturborotalita pseudopraebulloides</italic>, Sample A. Scale bars: <bold>(1)</bold>–<bold>(6b)</bold>, <bold>(7)</bold>–<bold>(9)</bold> <inline-formula><mml:math id="M26" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M27" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m; <bold>(6c)</bold> <inline-formula><mml:math id="M28" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 10 <inline-formula><mml:math id="M29" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m.</p></caption>
          <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f09.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
      <p id="d1e2279">Samples A–E contain abundant and diverse assemblages of planktonic
foraminifera, with a total of 40 species and 11 genera (Table 1). The
assemblages are characteristic of tropical–subtropical pelagic settings,
with common and abundant species of <italic>Globigerinatheka, Globoturborotalita</italic>, small (<inline-formula><mml:math id="M30" display="inline"><mml:mrow><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">250</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M31" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m)
<italic>Acarinina, Dentoglobigerina, Subbotina, Turborotalia</italic>, <italic>Hantkenina</italic> and <italic>Pseudohastigerina</italic>. <italic>Globigerina officinalis, Globorotaloides quadrocameratus, </italic>and<italic> Turborotalita quinqueloba</italic> are also present (Table 1, Figs. 4–10). Sample F was barren.</p>
      <p id="d1e2319">Preservation in Samples A to E ranges from very good to moderate. There is
some iron staining throughout. We use the zonal scheme of Berggren and
Pearson (2005) and Wade et al. (2011). The absence of the large muricate
taxa and the presence of <italic>Globigerinatheka semiinvoluta</italic> in Samples C, D and E indicate that the section
can be assigned to the planktonic foraminifera <italic>Globigerinatheka semiinvoluta</italic> HOZ (Zone E14; Berggren and
Pearson, 2005; Wade et al., 2011).</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F10" specific-use="star"><?xmltex \currentcnt{10}?><?xmltex \def\figurename{Figure}?><label>Figure 10</label><caption><p id="d1e2330">Stratigraphic section, planktonic foraminifera biostratigraphy
and range chart of species recorded in the Rashrashiyah Formation.
Taxa with earlier stratigraphic ranges than previously reported are in red;
later stratigraphic occurrences are in blue. Solid line indicates species
was found in the associated sample, and dashed lines are used when the species was not found in the sample but assumed to continue based on evidence
from elsewhere (e.g., Pearson et al., 2006; Wade et al., 2018a). Legend for
lithologies as in Fig. 3.</p></caption>
        <?xmltex \igopts{width=398.338583pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f10.png"/>

      </fig>

<?xmltex \hack{\newpage}?>
</sec>
<?pagebreak page155?><sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>Preservation</title>
      <p id="d1e2356">Four criteria were presented by Pearson and Burgess (2008) for
distinguishing foraminifera tests that are not significantly recrystallized:
(1) tests should be glassy or translucent in reflected light; (2) ultrafine
features such as spines (if initially possessed) should survive; (3) smooth
parts of the test such as the apertural lips, sutures, outer surface (in
some species) and inner surface (in most species) should appear smooth at
the submicron scale in high-resolution SEM images; and (4) in cross section
the submicron microgranular texture of the wall (if originally possessed)
should be clear when the test is broken.</p>
      <p id="d1e2359">Test preservation is very good in Sample A, specimens are glassy beneath the
light microscope and spines are preserved on <italic>Globigerinatheka</italic>, satisfying the criteria for
excellent preservation of Pearson and Burgess (2008). In Samples B and D,
the preservation is good, with specimens translucent under the light
microscope. Moderate preservation is recorded for Samples C and E, with some
infilling, encrustation and fragmentation (Figs. 4–9). Preservation is
superior to other sections in this region (e.g., Ramadan et al., 2021).</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>Biostratigraphy</title>
      <p id="d1e2373">The planktonic foraminiferal assemblage indicates that all samples are of
Priabonian age. No <italic>Morozovelloides</italic> or large <italic>Acarinina</italic> were found (except a singular dwarfed <italic>A. rohri</italic> in
Sample A), indicating that all samples are above the middle–late Eocene
turnover (MLET; Kamikuri and Wade, 2012; Wade et al., 2012). The extinction
of <italic>M. crassatus</italic> marks the base of the <italic>Globigerinatheka semiinvoluta </italic>HOZ Zone (Zone E14) (Berggren and Pearson, 2005;
Wade et al., 2011). The top (T) <italic>M. crassatus</italic> occurs within Chron C17n.3n and is
calibrated to 38.073 <inline-formula><mml:math id="M32" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.005 Ma at the Varignano section in Italy
(Luciani et al., 2020) based on the Pälike et al. (2006)
magnetochronology. This converts to 38.0 Ma on the Westerhold et al. (2014)
magnetochronology, which is the current standard used in the most recent
edition of the <italic>Geologic Time Scale</italic> (GTS2020; Speijer et al., 2020). The
entire section studied is therefore younger than 38 Ma.</p>
      <p id="d1e2405">We found a singular dwarfed specimen of <italic>Acarinina rohri</italic> in Sample A (Table 1). In the
western North Atlantic (ODP Site 1052) a reduction in specimen size of
<italic>Morozovelloides crassatus</italic> from 500 to 350 <inline-formula><mml:math id="M33" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m was recorded during the MLET, due to photosymbiont
bleaching and environmental stress (Wade et al., 2008; Wade and Olsson,
2009). Small <inline-formula><mml:math id="M34" display="inline"><mml:mrow><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">250</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M35" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m specimens of <italic>Acarinina</italic> consisting of <italic>A. collactea</italic>, <italic>A. echinata</italic> and <italic>A. medizzai</italic> are
found in all samples (Fig. 4). Small acarininids have been shown to range
beyond the MLET and into the Oligocene (Berggren et al., 2006; Wade and
Hernitz Kucenjak, 2018; Luciani et al., 2020), though they are not
consistently present in late Eocene samples from Jordan (Farouk et al.,
2015).</p>
      <p id="d1e2453"><italic>Globigerinatheka</italic> is common, with specimens of <italic>G. barri, G. index, G. korotkovi, G. kugleri, G. mexicana</italic>, <italic>G. semiinvoluta</italic>, and <italic>G. tropicalis</italic> (Premoli Silva et al., 2006).
<italic>Globigerinatheka semiinvoluta</italic> is present in samples C, D and E but absent from samples A and B. The Base (B) <italic>G. semiinvoluta </italic>is a secondary bioevent within Zone E14 but was previously used as a
primary bioevent to mark the base of Zone P15 (Berggren et al., 1995). It
has been calibrated to Chron C17n in several sections including ODP Site
1052<?pagebreak page156?> (Wade, 2004; Wade et al., 2012), Varignano (Luciani et al., 2020) and
Alano (Agnini et al., 2021). At the Varignano section, B <italic>G. semiinvoluta</italic> is calibrated to
Chron C17n.2n with an age of 37.665 <inline-formula><mml:math id="M36" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.006 Ma (timescale of Pälike
et al., 2006) (Luciani et al., 2020), which converts to 37.54 Ma on
Westerhold et al. (2014) magnetochronology (Fig. 11). The lack of <italic>G. semiinvoluta</italic> in
Samples A and B raises the possibility that these samples could be from the
short stratigraphic interval between the T <italic>M. crassatus</italic> and the B <italic>G. semiinvoluta</italic>. This interval was
designated the <italic>Turborotalia pseudoampliapertura</italic> Zone by Haggag (1990) and has been recognized in several
sections worldwide (Canudo and Molina, 1992; Wade, 2004; Agnini et al.,
2011; Strougo et al., 2013). The B <italic>G. semiinvoluta</italic> may be diachronous, as the duration of
the interval between T <italic>M. crassatus</italic> and B <italic>G. semiinvoluta</italic> is only 20 kyr at ODP Site 1052 (Wade, 2004;
Wade et al., 2012), 330 kyr at Alano and 408 kyr at the Varignano section
(Luciani et al., 2020). Note, however, the discrepancy could be due to the
criteria used to separate <italic>G. semiinvoluta</italic> from its ancestor <italic>G. mexicana</italic> (Premoli Silva et al., 2006).
Thus the bioevent may be isochronous, but independent workers have not been
unified in the discrimination of the first true <inline-formula><mml:math id="M37" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>.</mml:mo></mml:mrow></mml:math></inline-formula> <italic>semiinvoluta</italic> morphospecies.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F11"><?xmltex \currentcnt{11}?><?xmltex \def\figurename{Figure}?><label>Figure 11</label><caption><p id="d1e2529">Section log of the upper Eocene Rashrashiyah Formation with
planktonic foraminifera and calcareous nannofossil zonal schemes and
bioevents. Drawing of <italic>G. semiinvoluta</italic> from Wade (2004). Legend for lithologies as in Fig. 3.</p></caption>
          <?xmltex \igopts{width=236.157874pt}?><graphic xlink:href="https://jm.copernicus.org/articles/40/145/2021/jm-40-145-2021-f11.png"/>

        </fig>

      <p id="d1e2541">T <italic>Planorotalites capdevilensis</italic> is slightly older than B <italic>G. semiinvoluta</italic> in the Italian sections of Alano (Agnini et
al., 2011) and Varignano (Luciani et al., 2020). Wade (2004) found a longer
range of <italic>P. capdevilensis</italic> from ODP Site 1052, western North Atlantic. <italic>P. capdevilensis</italic> was not found in this
study of the Rashrashiyah Formation.</p>
      <p id="d1e2556">The characteristics of the assemblage are very consistent with stratigraphic
equivalent sections studied in the Adriatic, Egypt and Armenia (Wade et al.,
2012; Strougo et al., 2013; Cotton et al., 2017; Ramadan et al., 2021;
Salama et al., 2021). However, despite the high diversity, no <italic>Catapsydrax</italic> are present.
This contrasts Adriatic cores where <italic>Catapsydrax</italic> increase in abundance at the base
Priabonian (Wade et al., 2012). We record a high diversity of species
belonging to <italic>Globoturborotalita</italic> (Figs. 8, 9 and 10), and <italic>G. pseudopraebulloides</italic> is unusually abundant (Fig. 9).</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>Stratigraphic ranges</title>
      <p id="d1e2579">The stratigraphic distribution of the recorded species is shown in Fig. 10. The stratigraphic ranges of many of the non-marker species are different
to published schemes (e.g., Pearson et al., 2006; Aze et al., 2011; Wade et
al., 2018a). Many species thought to have evolved in Zone E16 or Zone O1 are
found at this section in Zone E14, suggesting their stratigraphic range
needs revision. For example, <italic>Globoturborotalita barbula, Globoturborotalita pseudopraebulloides</italic>, <italic>Globoturborotalita paracancellata</italic>, <italic>Globoturborotalita cancellata</italic>, <italic>Globoturborotalita gnaucki</italic> and <italic>Subbotina projecta</italic> (Figs. 5, 8 and 9) were all thought to
evolve between upper Eocene Zone E15 and lower Oligocene Zone O2
(Spezzaferri et al., 2018; Wade et al., 2018b, c). However, they are all
present in Zone E14, with some species, for instance, <italic>Globoturborotalita pseudopraebulloides</italic> in high abundance (Fig. 9). The
discrepancy between first occurrence in the published literature and this
study is most likely due to the excellent preservation of this section
coupled with the lack of recent taxonomic investigations on the late Eocene.
Within the combined nannofossil and planktonic foraminifera assemblages, we
do not see any evidence for reworking in these sediments as an explanation
for the lower occurrences of certain species compared to their previously
published stratigraphic ranges.</p>
      <p id="d1e2601"><italic>Dentoglobigerina</italic> is a diverse genus that evolved in the middle Eocene. We find and
illustrate (Fig. 5.6 and 5.7) two specimens that we could not confidently
place in any of the previously described species: we refer to these as
<italic>Dentoglobigerina</italic> sp. 1 and <italic>Dentoglobigerina</italic> sp. 2, pending further investigations (see Appendix A for
taxonomic notes). Many of the stratigraphic ranges presented in Pearson et
al. (2006) have already been extended to older levels by Wade et al. (2018a)
and Fayolle and Wade (2021). <italic>Dentoglobigerina eotripartita</italic> is present from the base sample (Sample A),
confirming the stratigraphic range suggested in Wade et al. (2018a).
<italic>Dentoglobigerina taci</italic> was thought to be confined to the Eocene–Oligocene boundary interval (Zone E16 to Zone O1) (Pearson and Wade, 2015; Wade et al., 2018a). Here we find
<italic>Dentoglobigerina taci</italic> in Zone E14 (Figs. 5 and 10), extending its evolution to several million
years earlier. We find a singular rare occurrence of <italic>D. tripartita</italic> in Sample C (Fig. 5.5).</p>
      <p id="d1e2625">Globigerinathekids are abundant in the samples. Premoli Silva et al. (2006)
state that the extinction of <italic>G. barri</italic> occurs towards the end of Zone E14. Here we
find <italic>G. barri</italic> in Samples A, B and C (Fig. 6), but not in the younger part of<?pagebreak page157?> the
section, potentially constraining the extinction of <italic>G. barri</italic> to early Zone E14. Our
extinction horizon for <italic>G. barri</italic> is consistent with results from coeval Egyptian
sections (Strougo et al., 2013). In Premoli Silva et al. (2006) the
extinction of <italic>Globigerinatheka kugleri </italic>is given as Zone E13, but we find this species ranging higher
(Zone E14).</p>
      <p id="d1e2643"><italic>Globoturborotalita</italic> is a long-ranging genus, from the Eocene to the present. Many species have
been described, though their ranges, in general, are poorly constrained.
<italic>Globoturborotalita barbula</italic> was previously only known from the Eocene–Oligocene boundary interval
(Pearson and Wade, 2015; Spezzaferri et al., 2018). We find specimens in
Samples A and B, suggesting this species evolved earlier than previously
thought (Pearson and Wade, 2015; Spezzaferri et al., 2018). The
stratigraphic range of <italic>G. cancellata</italic> is not well constrained, and until Spezzaferri et
al. (2018) it had not been recorded outside of the Oligocene <italic>Globorotalia opima opima</italic> Zone, from
which it was described. Spezzaferri et al. (2018) found and illustrated
specimens from Zone O1 and suggested a questionable range from Zone E16 to
Zone O5. We find and illustrate specimens from Zone E14 (Fig. 8), indicating
that this species has a much longer stratigraphic range than previously
suggested. <italic>G. paracancellata</italic> was described from the upper Oligocene, with recorded specimens
occurring from the upper Eocene Zone E16 (Spezzaferri et al., 2018). Here we
extend the stratigraphic range of <italic>G. paracancellata</italic> with specimens illustrated from Zone E14
(Fig. 9). <italic>G. gnaucki </italic>is abundant in Samples B, C and D (Table 1, Fig. 8). This species
was thought to evolve in Zone E15 (Spezzaferri et al., 2018), but this study
suggests it evolved in Zone E14 or older (Fig. 10). We find specimens that
we refer to as <italic>Globoturborotalita</italic> cf. <italic>G. labiacrassata</italic> (Fig. 9). These have a high-arched umbilical aperture
and lobate profile but lack the thick rim boarding the aperture that is
characteristic of this species. Our specimens are in Zone E14 and thus much
older than the first appearance of <italic>G. labiacrassata</italic> (Zone O2) suggested by Spezzaferri et
al. (2018). Further investigations are required to determine if these forms
are a new species.</p>
      <p id="d1e2677">In Pearson et al. (2006) the evolution of <italic>Pseudohastigerina naguewichiensis</italic> from <italic>P. micra</italic> occurs at the base of Zone
E15. We find and illustrate <italic>P. naguewichiensis</italic> in Zone E14 (Fig. 4). Our earlier evolution of
this species is in agreement with Cotton et al. (2017), though an older
range (Zone E13) is suggested by Strougo et al. (2013). The oldest
previously recorded specimens of <italic>Subbotina projecta</italic> are from upper Eocene Zone E16. Here we
document and illustrate specimens from Zone E14 (Table 1, Figs. 5 and 10).
The occurrence of <italic>T. quinqueloba</italic> in Sample D (Figs. 8 and 10) confirms the stratigraphic
range recorded in Pearson and Kučera (2018).</p>
      <p id="d1e2695">Despite the relatively good preservation and the high diversity of species
recorded from the Rashrashiyah Formation (Table 1, Figs. 4–10), there are
some species that were expected to be present, based on previous range chart
compilations (e.g., Pearson et al., 2006; Aze et al., 2011), that were not
found. These include <italic>Paragloborotalia griffinoides</italic> and <italic>P. nana</italic>, <italic>Chiloguembelina</italic> <italic>ototara</italic>, <italic>C. cubensis</italic>, and species of the genus <italic>Catapsydrax</italic>. However, we
note that these species are recorded in upper Eocene sections from Egypt
(Ramadan et al., 2021). We suspect that the absence of these taxa in the
Rashrashiyah Formation is due to the environment, which is indicative of
warm and oligotrophic conditions.</p>
      <p id="d1e2717">The amendment of planktonic foraminifera stratigraphic ranges has
ramifications for the phylogenies and tropical–subtropical diversity charts.
The range charts presented in Pearson et al. (2006), Wade et al. (2018a),
incorporated into Time-Scale Creator (Fordham et al., 2018), and the Mikrotax
online portal (Huber et al., 2016) will require revision. Many species
evolved earlier than previously thought, particularly within the
<italic>Globoturborotalita</italic> genus. The extensions of the range of <italic>Pseudohastigerina</italic> <italic>naguewichiensis</italic> and <italic>Turborotalita quinqueloba</italic> were already suggested in
Cotton et al. (2017) and confirmed here. Our study implies that the late
Eocene tropical–subtropical diversity is higher than previously suggested in
compilations (Ezard et al., 2011; Fraass et al., 2015; Lowery et al., 2020).
Many of the species found in this study of the late Eocene extend through
the Eocene–Oligocene transition and into the Oligocene, suggesting that the
rate of turnover at the Eocene–Oligocene transition is not as large as
previously thought and requires re-investigation.</p>
</sec>
<sec id="Ch1.S4.SS4">
  <label>4.4</label><title>Integrated calcareous biostratigraphy</title>
      <p id="d1e2740">The calcareous nannofossil assemblages were studied by Aljahdali et al. (2020) indicating that the section corresponds to Zone CNE17 of Agnini et
al. (2014) and Zone NP18 of Martini (1971). The section is Priabonian age
(upper Eocene), as indicated by the presence of <italic>Chiasmolithus oamaruensis.</italic> The T <italic>C. grandis</italic> between Samples B
and C allows identification of the base Zone CP15 of Okada and Bukry (1980).
The base common (Bc) of <italic>Cribrocentrum erbae</italic> also occurs between Samples B and C. Throughout the
section <italic>C. erbae</italic> increases in abundance consist with Zone CNE17 of Agnini et al. (2014). A single specimen of <italic>Isthmolithus recurvus</italic> was documented in Sample E (Aljahdali et al.,
2020); however, this is not used in the biostratigraphic interpretation. The
planktonic foraminifera biostratigraphy is consistent with the calcareous
nannofossil biostratigraphy and can also be compared to other
integrated calcareous biostratigraphic studies (e.g., Strougo et al., 2013;
Farouk et al., 2015; Cotton et al., 2017; Luciani et al., 2020; Agnini et
al., 2021).</p>
      <p id="d1e2758">The base of the Priabonian Global Stratotype Section and Point (GSSP) was
recently defined by Agnini et al. (2021) at the Alano di Piave section
(north-eastern Italy). The Bartonian–Priabonian boundary is placed at the
prominent 14–16 cm crystal tuff layer known as the “Tiziano bed” at 63.57 m.
With our current 1 m sampling resolution and the three bioevents between
Samples B and C, we are unable to confidently calculate sedimentation rates,
but it would appear<?pagebreak page158?> that the base of the section is either at, or very close
to, the Bartonian–Priabonian boundary.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d1e2771">The planktonic foraminiferal biostratigraphy, integrated with calcareous
nannofossil biostratigraphy, provides a robust stratigraphic framework for
the Rashrashiyah Formation, indicating that the section is Priabonian (upper
Eocene) in age. Planktonic foraminifera assemblages are diverse and
extremely well-preserved. Our study reveals differing stratigraphic ranges
to the ones established in the literature. Higher-resolution sampling will
allow the horizons for Base <italic>G. semiinvoluta</italic>, Top <italic>C. grandis</italic> and Base Common <italic>C. erbae</italic> to be differentiated.
Our study implies that the late Eocene tropical–subtropical diversity is
likely to be higher than previously suggested in data compilations.</p>
</sec>

      
      </body>
    <back><app-group>

<app id="App1.Ch1.S1">
  <?xmltex \currentcnt{A}?><label>Appendix A</label><title>Taxonomic list of species in this study</title>
      <p id="d1e2794"><table-wrap id="Taba" position="anchor"><oasis:table><oasis:tgroup cols="1">
     <oasis:colspec colnum="1" colname="col1" align="justify" colwidth="8cm"/>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Acarinina collactea</italic> (Finlay).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Acarinina echinata</italic> (Bolli), Fig. 4.1.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Acarinina medizzai</italic> (Toumarkine and Bolli), Fig. 4.2–4.6.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Acarinina rohri </italic>(Brönnimann and Bermúdez).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina eotripartita</italic> Pearson, Wade, and <?xmltex \hack{\hfill\break}?>Olsson, Fig. 5.1.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina galavisi</italic> (Bermúdez), Fig. 5.2.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina pseudovenezuelana</italic> <?xmltex \hack{\hfill\break}?>(Blow and Banner).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina taci</italic> Pearson and Wade, Fig. 5.3 and 5.4.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina tripartita</italic> (Koch), Fig. 5.5.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina </italic>sp. 1, Fig. 5.7. This specimen has a dentoglobigerinid wall texture, a compressed final chamber and pronounced tooth.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Dentoglobigerina </italic>sp. 2, Fig. 5.6. This specimen has incised umbilical sutures and a compressed final chamber. In umbilical view this specimen bears a close morphological resemblance to the drawing of <italic>Globigerinatheka index</italic> by Postuma (1971). However, due to the wall texture and lack of supplementary apertures, we consider this specimen to belong within <italic>Dentoglobigerina</italic>, pending further investigations.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerina officinalis </italic>Subbotina.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka barri</italic> Brönnimann, Fig. 6.3 and 6.4.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka index </italic>(Finlay), Fig. 6.5 and 6.6.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka korotkovi</italic> (Keller).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka kugleri</italic> (Bolli, Loeblich, and Tappan), Fig. 6.7 and 6.8. <italic>Globigerinatheka mexicana</italic> (Cushman), Fig. 6.9 and  6.10.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka semiinvoluta</italic> (Keijzer), Fig. 7.4 and <?xmltex \hack{\hfill\break}?>7.5.</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>
        <?xmltex \hack{\newpage}?><table-wrap id="Tabb" position="anchor"><oasis:table><oasis:tgroup cols="1">
     <oasis:colspec colnum="1" colname="col1" align="justify" colwidth="8cm"/>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globigerinatheka tropicalis</italic> (Blow and Banner), <?xmltex \hack{\hfill\break}?>Fig. 7.1–7.3.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globorotaloides quadrocameratus</italic> Olsson, Pearson, and Huber, Fig. 7.6.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita barbula</italic> Pearson and Wade, Fig. 8.1 and 8.2.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita cancellata</italic> (Pessagno), Fig. 8.3–8.5.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita gnaucki</italic> (Blow and Banner), Fig. 8.6.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita </italic>cf.<italic> G. labiacrassata</italic> (Jenkins), <?xmltex \hack{\hfill\break}?>Fig. 9.1 and 9.2. These specimens have a high-arched umbilical aperture and lobate profile but lack the thick rim boarding the aperture that is characteristic of <italic>G. labiacrassata</italic>.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita ouachitaensis</italic> (Howe and Wallace), Fig. 8.7 and 8.8.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita paracancellata</italic> Olsson and <?xmltex \hack{\hfill\break}?>Hemleben, Fig. 9.3 and 9.4.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Globoturborotalita pseudopraebulloides</italic> Olsson and <?xmltex \hack{\hfill\break}?>Hemleben, Fig. 9.5–9.9.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Hantkenina alabamensis </italic>Cushman.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Hantkenina primitiva </italic>Cushman and Jarvis, Fig. 6.2.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Pseudohastigerina micra</italic> (Cole), Fig. 4.9–4.12.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Pseudohastigerina naguewichiensis </italic>(Myatliuk), Fig. 4.7 and 4.8.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Subbotina corpulenta </italic>(Subbotina), Fig. 6.1.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Subbotina linaperta</italic> (Finlay).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Subbotina projecta</italic> Olsson, Pearson, and Wade, Fig. 5.8.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Subbotina utilisindex</italic> Jenkins and Orr, Fig. 5.9 and 5.10.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Subbotina yeguanensis</italic> (Weinzierl and Applin).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalia ampliapertura </italic>(Bolli), Fig. 7.7.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalia cerroazulensis </italic>(Cole), Fig. 7.8–7.10.</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalia cunialensis </italic>(Toumarkine and Bolli).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalia increbescens</italic> (Bandy).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalia pomeroli </italic>(Toumarkine and Bolli).</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><italic>Turborotalita quinqueloba</italic> (Natland), Fig. 8.9.</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap></p>
</app>
  </app-group><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d1e3184">The data generated in this study are included within the paper and in Table 1. Imaged specimens are deposited at the Natural History Museum, London, UK.</p>
  </notes><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d1e3190">BW conducted the analyses and wrote the paper. MA conceived the project and
the prepared lithographic logs. MA, YM, AM, SA and IZ conducted fieldwork,
sampled the studied section and provided comments to manuscript text and
figures.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d1e3196">The authors declare that they have no conflict of interest.</p>
  </notes><notes notes-type="disclaimer"><title>Disclaimer</title>

      <p id="d1e3202">Publisher’s note: Copernicus Publications remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p>
  </notes><ack><title>Acknowledgements</title><p id="d1e3208">We are extremely grateful to Natalie Cheng, who conducted the light and SEM
imaging, prepared the plates and assisted in taxonomic discussion, and to
Marcin Latas, who assisted with sample preparation and the map. Comments on
an earlier version of the manuscript were provided by Paul Pearson, Florent Fayolle and Alessio Fabbrini. We thank the CEO of the Saudi Geological
Survey Eng. Abdullah M. Al-Shamrani and vice president Saleh Al-Sefry, Nasser Aljahdali and Wadee Kashghari for supporting fieldwork, permission to use
SGS labs and equipment for this work. We thank Helen Coxall and an anonymous reviewer for their constructive
suggestions. This paper was edited by Kirsty Edgar, who provided additional
insights and comments that improved the manuscript.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d1e3213">Bridget S. Wade was supported by UK Natural Environment Research Council (NERC) reference
number NE/G014817.</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d1e3219">This paper was edited by Kirsty Edgar and reviewed by Helen Coxall and one anonymous referee.</p>
  </notes><ref-list>
    <title>References</title>

      <ref id="bib1.bib1"><label>1</label><?label 1?><mixed-citation>
Agnini, C., Fornaciari, E., Giusberti, L., Grandesso, P., Lanci, L., Luciani, V., Muttoni, G., Palike, H., Rio, D., Spofforth, D. J. A., and Stefani, C.:  Integrated biomagnetostratigraphy of the Alano section (NE Italy): A proposal for defining the middle-late Eocene boundary, Geol. Soc. Am. B., 123, 841–872, 2011.</mixed-citation></ref>
      <ref id="bib1.bib2"><label>2</label><?label 1?><mixed-citation>
Agnini, C., Fornaciari, E., Raffi, I., Catanzariti, R., Pälike, H.,
Backman, J., and Rio, D.: Biozonation and biochronology of Paleogene calcareous
nannofossils from low and middle latitudes, Newsl. Stratigr., 47,
131–181, 2014.</mixed-citation></ref>
      <ref id="bib1.bib3"><label>3</label><?label 1?><mixed-citation>
Agnini, C., Backman, J., Boscolo-Galazzo, F., Condon, D. J., Fornaciari, E.,
Galeotti, S., Giusberti, L., Grandesso, P., Lanci, L., Luciani, V., Monechi,
S., Muttoni, G., Pälike, H., Pampaloni, M. L., Papazzoni, C. A., Pearson,
P. N., Pignatti, J., Premoli Silva, I., Raffi, I., Rio, D., Rook, L., Sahy,
D., Spofforth, D. J. A., Stefani, C., and Wade, B. S.: Proposal for the Global
Boundary Stratotype Section and Point (GSSP) for the Priabonian Stage
(Eocene) at the Alano section (Italy), Episodes, 44, 151–173, 2021.</mixed-citation></ref>
      <ref id="bib1.bib4"><label>4</label><?label 1?><mixed-citation>Aljahdali, M. H., Elhag, M., Mufrreh, Y., Memesh, A., AlSoubhi, S., and
Zalmout, I. S.: Upper Eocene calcareous nannofossil biostratigraphy: a new
preliminary Priabonian record from northern Saudi Arabia, Appl. Ecol.
Environ. Res., 18, 5607–5625, <ext-link xlink:href="https://doi.org/10.15666/aeer/1804_56075625" ext-link-type="DOI">10.15666/aeer/1804_56075625</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib5"><label>5</label><?label 1?><mixed-citation>Al-Rawi, M. M.: Petroleum systems in Jordan, GEO ExPro, 11, 1, available at: <uri>https://archives.datapages.com/data/geo-expro-magazine/011/011001/pdfs/22.htm</uri> (last access: 26 September 2021), 2014.</mixed-citation></ref>
      <ref id="bib1.bib6"><label>6</label><?label 1?><mixed-citation>
Aze, T., Ezard, T. H. G., Purvis, A., Coxall, H. K., Stewart, D. R. M., Wade, B. S.,
and Pearson, P. N.: A phylogeny of Cenozoic macroperforate planktonic
foraminifera from fossil data, Biol. Rev., 86, 900–927, 2011.</mixed-citation></ref>
      <ref id="bib1.bib7"><label>7</label><?label 1?><mixed-citation>
Berggren, W. A. and Pearson, P. N.: A revised tropical to subtropical planktonic
foraminiferal zonation of the Eocene and Oligocene, J. Foramin.
Res., 35, 279–298, 2005.</mixed-citation></ref>
      <ref id="bib1.bib8"><label>8</label><?label 1?><mixed-citation>
Berggren, W. A., Kent, D. V., Swisher III, C. C., and Aubry, M.-P.: A Revised
Cenozoic Geochronology and Chronostratigraphy, in:
Geochronology, Time Scales and Global Stratigraphic Correlation, edited by: Berggren, W. A., Kent, D. V., Aubry, M.-P., and
Hardenbol, J., SEPM
Society for Sedimentary Geology, Tulsa, OK, USA, 1995.</mixed-citation></ref>
      <ref id="bib1.bib9"><label>9</label><?label 1?><mixed-citation>Berggren, W. A., Pearson, P. N., Huber, B. T., and Wade, B. S.: Taxonomy,
Biostratigraphy and Phylogeny of Eocene <italic>Acarinina</italic>,  in: Atlas of Eocene
Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K.,
Huber, B. T., Hemleben, C., and Berggren, W. A., Cushman Foundation Special Publication, 41,
257–326, 2006.</mixed-citation></ref>
      <ref id="bib1.bib10"><label>10</label><?label 1?><mixed-citation>
Bown, P. R., Dunkley Jones, T., Lees, J. A., Randell, R. D., Mizzi, J. A.,
Pearson, P. N., Coxall, H. K., Young, J. R., Nicholas, C. J., Karega, A.,
Singano, J., and Wade, B. S.: A Paleogene calcareous microfossil
Konservat-Lagerstätte from the Kilwa Group of coastal Tanzania,
Geol. Soc. Am. Bull., 120, 3–12, 2008.</mixed-citation></ref>
      <ref id="bib1.bib11"><label>11</label><?label 1?><mixed-citation>
Canudo, J. I. and Molina, E.: Bioestratigrafía con foraminíferos
planctónicos del Paleógeno del Pirineo. in h. Luterbacher (edn.),
Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 186,
97–135, 1992.</mixed-citation></ref>
      <ref id="bib1.bib12"><label>12</label><?label 1?><mixed-citation>Cotton, L. J., Zakrevskaya, E. Y., van der Boon, A., Asatryan, G., Hayrapetyan,
F., Israyelyan, A., Krijgsman, W., Less, G., Monechi, S., Papazzoni, C. A.,
Pearson, P. N., Razumovskiy, A., Renema, W., Shcherbinina, E., and Wade,
B. S.: Integrated stratigraphy of the Priabonian (upper Eocene) Urtsadzor
section, Armenia, Newsl. Stratigr., 50, 269–295, <ext-link xlink:href="https://doi.org/10.1127/nos/2016/0313" ext-link-type="DOI">10.1127/nos/2016/0313</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib13"><label>13</label><?label 1?><mixed-citation>Dunkley Jones, T., Bown, P. R., Pearson, P. N., Wade, B. S., Coxall, H. K., and
Lear, C. H.: Major shifts in calcareous phytoplankton assemblages through the
Eocene-Oligocene transition of Tanzania and their implications for
low-latitude primary production, Paleoceanography, 23, PA4204,
<ext-link xlink:href="https://doi.org/10.1029/2008PA001640" ext-link-type="DOI">10.1029/2008PA001640</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib14"><label>14</label><?label 1?><mixed-citation>
Ezard, T. H. G., Aze, T., Pearson, P. N., and Purvis, A.: Interplay between
changing climate and species' ecology drives macroevolutionary dynamics,
Science, 332, 349–351, 2011.</mixed-citation></ref>
      <ref id="bib1.bib15"><label>15</label><?label 1?><mixed-citation>
Farouk, S., Faris, M., Ahmad, F., and Powell, J. H.: New microplanktonic
biostratigraphy and depositional sequences across the Middle-Late Eocene and
Oligocene boundaries in eastern Jordan, GeoArabia – Middle East Petroleum
Geosciences, 20, 145–172, 2015.</mixed-citation></ref>
      <ref id="bib1.bib16"><label>16</label><?label 1?><mixed-citation>Fayolle, F. and Wade, B. S.: The evolution of Eocene planktonic foraminifera
<italic>Dentoglobigerina</italic>, J. Syst. Palaeontol., 19, 333–376, 2021.</mixed-citation></ref>
      <ref id="bib1.bib17"><label>17</label><?label 1?><mixed-citation>
Firth, J. V., Eldrett, J. S., Harding, I. C., Coxall, H. K., and Wade, B. S.:
Integrated biomagnetochronology for the Palaeogene of ODP Hole 647A:
implications for correlating palaeoceanographic events from high to low
latitudes, in: Magnetic Methods and the Timing of Geological Processes, edited by: Jovane, L., Herrero-Bervera, E., Hinnov, L. A., and Housen,
B. A.,
Geological Society, London, Special Publications, 373, 29–78, 2013.</mixed-citation></ref>
      <ref id="bib1.bib18"><label>18</label><?label 1?><mixed-citation>Fordham, B. G., Aze, T., Haller, C., Zehady, A. K., Pearson, P. N., Ogg, J. G.,
and Wade, B. S.: Future-proofing the Cenozoic macroperforate planktonic
foraminifera phylogeny of Aze &amp; others (2011), PLoS ONE, 13, e0204625,
<ext-link xlink:href="https://doi.org/10.1371/journal.pone.0204625" ext-link-type="DOI">10.1371/journal.pone.0204625</ext-link>, 2018.</mixed-citation></ref>
      <?pagebreak page160?><ref id="bib1.bib19"><label>19</label><?label 1?><mixed-citation>Fraass, A. J., Kelly, D. C., and Peters, S. E.: Macroevolutionary history of the
planktic foraminifera, Annu. Rev. Earth Planet. Sc., 43,
139–166, <ext-link xlink:href="https://doi.org/10.1146/annurev-earth-060614-105059" ext-link-type="DOI">10.1146/annurev-earth-060614-105059</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib20"><label>20</label><?label 1?><mixed-citation>
Guiraud, R., Issawi, B., and Bosworth, W.: Phanerozoic history of Egypt and
surrounding areas, Peri-Tethys Memoir, 6, 469–509, 2001.</mixed-citation></ref>
      <ref id="bib1.bib21"><label>21</label><?label 1?><mixed-citation>Haggag, M. A.: <italic>Globigerina pseudoampliapertura</italic> Zone, a new late Eocene planktonic foraminiferal zone (Fayoum
area, Egypt), Neues Jahrbuch für Geologie und Paläontologie,
Monatshefte, Stuttgart, 295–307, 1990.</mixed-citation></ref>
      <ref id="bib1.bib22"><label>22</label><?label 1?><mixed-citation>
Halawani, M.: Stratigraphic column for the Phanerozoic rocks of Saudi
Arabia. A compilation and synthesis with comments – Saudi Geological
Survey Technical Reports, Kingdom of Saudi Arabia SGS-TR-2001-3 2001,
2001.</mixed-citation></ref>
      <ref id="bib1.bib23"><label>23</label><?label 1?><mixed-citation>
Hemleben, C. and Olsson, R. K.: Wall textures of Eocene planktonic
foraminifera, in: Atlas of Eocene Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and
Berggren, W. A., Cushman
Foundation Special Publication, 41, 47–66, Allen Press, Lawrence, KS, USA, 2006.</mixed-citation></ref>
      <ref id="bib1.bib24"><label>24</label><?label 1?><mixed-citation>
Huber, B. T., Petrizzo, M. R., Young, J., Falzoni, F., Gilardoni, S., Bown,
P. R., and Wade, B. S.: Pforams@mikrotax: A new online taxonomic database for
planktonic foraminifera, Micropalaeontology, 62, 429–438, 2016.</mixed-citation></ref>
      <ref id="bib1.bib25"><label>25</label><?label 1?><mixed-citation>
Kamikuri, S. and Wade, B. S.: Radiolarian magnetobiochronology and faunal
turnover across the middle/late Eocene boundary at Ocean Drilling Program
Site 1052 in the western North Atlantic Ocean, Mar. Micropaleontol.,
88–89, 41–53, 2012.</mixed-citation></ref>
      <ref id="bib1.bib26"><label>26</label><?label 1?><mixed-citation>
Lear, C., Bailey, T. R., Pearson, P. N., Coxall, H. K., and Rosenthal, Y.:
Cooling and ice growth across the Eocene – Oligocene transition, Geology,
36, 251–254, 2008.</mixed-citation></ref>
      <ref id="bib1.bib27"><label>27</label><?label 1?><mixed-citation>
Lowery, C. M., Bown, P. R., Fraass, A. J., and Hull, P. M.: Ecological response
of plankton to environmental change: thresholds for extinction, Annu.
Rev. Earth Planet. Sc., 48, 403–429, 2020.</mixed-citation></ref>
      <ref id="bib1.bib28"><label>28</label><?label 1?><mixed-citation>
Luciani, V., Fornaciari, E., Papazzoni, C. A., Dallanave, E., Giusberti, L.,
Stefani, C., and Amante, E.: Integrated stratigraphy at the
Bartonian–Priabonian transition: correlation between shallow benthic and
calcareous plankton zones (Varignano section, northern Italy), Geol.
Soc. Am. Bull., 132, 495–520, 2020.</mixed-citation></ref>
      <ref id="bib1.bib29"><label>29</label><?label 1?><mixed-citation>
Martini, E.: Standard Tertiary and Quaternary calcareous nannoplankton
zonation. Proc. II Planktonic Conference, Roma, 1970, Tecnoscienza, Roma,
1971.</mixed-citation></ref>
      <ref id="bib1.bib30"><label>30</label><?label 1?><mixed-citation>
Meissner, C. R., Griffin Jr., M. B., Riddler, G. P., Marcel Van Eck,
Aspinall, N. C., Farasani, A. M., and Dini, S. M.: Preliminary Geologic Map of
the Thaniyat Turayf Quadrangle, Sheet 29c, Kingdom of Saudi Arabia –
Department of the Interior, US Geological Survey, Washington, DC, 1990.</mixed-citation></ref>
      <ref id="bib1.bib31"><label>31</label><?label 1?><mixed-citation>
Miller, K. G., Browning, J. V., Aubry, M.-P., Wade, B. S., Katz, M. E., Kulpecz,
A. A., and Wright, J. D.: Eocene-Oligocene global climate and sea-level
changes: St. Stephens Quarry, Alabama, Geol. Soc. Am. Bull.,
120, 34–53, 2008.</mixed-citation></ref>
      <ref id="bib1.bib32"><label>32</label><?label 1?><mixed-citation>
Moore, T. C. and Kamikuri, S.: Data report: Radiolarian stratigraphy across
the Eocene/Oligocene boundary in the equatorial Pacific, Sites 1218, U1333,
and U1334, In: Proceedings IODP, vol. 320/321, Pälike et
al., Integr. Ocean Drill. Program Management International, Inc., Tokyo,
2012.</mixed-citation></ref>
      <ref id="bib1.bib33"><label>33</label><?label 1?><mixed-citation>
Norris, R. D., Wilson, P. A., Blum, P., and the Expedition 342 Scientists:
Proc. IODP, 342: College Station, TX (Integrated Ocean Drilling Program), College Station, TX, USA,
2014.</mixed-citation></ref>
      <ref id="bib1.bib34"><label>34</label><?label 1?><mixed-citation>
Okada, H. and Bukry, D.: Supplementary modification and introduction of code
numbers to the low latitude coccolith biostratigraphic zonation (Bukry,
1973; 1975), Mar. Micropaleontol., 5, 321–325, 1980.</mixed-citation></ref>
      <ref id="bib1.bib35"><label>35</label><?label 1?><mixed-citation>
Pälike, H., Norris, R. D., Herrle, J. O., Wilson, P. A., Coxall, H. K.,
Lear, C. H., Shackleton, N. J., Tripati, A. K., and Wade, B. S.: The heartbeat of
the Oligocene climate system, Science, 314, 1894–1898, 2006.</mixed-citation></ref>
      <ref id="bib1.bib36"><label>36</label><?label 1?><mixed-citation>
Pearson, P. N. and Burgess, C. E.: Foraminifer test preservation and
diagenesis: comparison of high latitude Eocene sites, Geol. Soc.
Lond. Spec. Publ., 303, 59–72, 2008.</mixed-citation></ref>
      <ref id="bib1.bib37"><label>37</label><?label 1?><mixed-citation>Pearson, P. N. and Kučera, M.: Taxonomy, biostratigraphy, and phylogeny
of Oligocene <italic>Turborotalita</italic>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and
Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46, 385–392,
2018.</mixed-citation></ref>
      <ref id="bib1.bib38"><label>38</label><?label 1?><mixed-citation>
Pearson, P. N. and Wade, B. S.: Systematic taxonomy of exceptionally
well-preserved planktonic foraminifera from the Eocene/Oligocene boundary of
Tanzania, Cushman Foundation for Foraminiferal Research Special Publication,
No. 45, 1–85, Allen Press, Lawrence, KS, USA,  2015.</mixed-citation></ref>
      <ref id="bib1.bib39"><label>39</label><?label 1?><mixed-citation>
Pearson, P. N., Ditchfield, P. W., Singano, J., Harcourt-Brown, K. G.,
Nicholas, C. J., Olsson, R. N., Shackleton, N. J., and Hall, M. A.: Warm
tropical sea surface temperatures in the Late Cretaceous and Eocene epochs,
Nature, 413, 481–487, 2001.</mixed-citation></ref>
      <ref id="bib1.bib40"><label>40</label><?label 1?><mixed-citation>
Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and Berggren, W. A.:
Atlas of Eocene planktonic foraminifera, Cushman Foundation for
Foraminiferal Research Special Publication, 41, 513 pp., Allen Press, Lawrence, KS, USA, 2006.</mixed-citation></ref>
      <ref id="bib1.bib41"><label>41</label><?label 1?><mixed-citation>
Pearson, P. N., van Dongen, B. E., Nicholas, C. J., Pancost, R. D., Schouten,
S., Singano, J. M., and Wade, B. S.: Stable warm tropical climate through the
Eocene epoch, Geology, 35, 211–214, 2007.</mixed-citation></ref>
      <ref id="bib1.bib42"><label>42</label><?label 1?><mixed-citation>
Pearson, P. N., McMillan, I. K., Wade, B. S., Dunkley Jones, T., Coxall, H. K.,
Bown, P. R., and Lear, C. H.: Extinction and environmental change across the
Eocene-Oligocene boundary in Tanzania, Geology, 36, 179–182, 2008.</mixed-citation></ref>
      <ref id="bib1.bib43"><label>43</label><?label 1?><mixed-citation>
Pearson, P. N., Foster, G. L., and Wade, B. S.: Atmospheric carbon dioxide
through the Eocene-Oligocene climate transition, Nature, 461, 1110–1113,
2009.</mixed-citation></ref>
      <ref id="bib1.bib44"><label>44</label><?label 1?><mixed-citation>
Postuma, J. A.: Manual of planktonic foraminifera, Elsevier for Shell Group,
The Hague, 406 pp., 1971.</mixed-citation></ref>
      <ref id="bib1.bib45"><label>45</label><?label 1?><mixed-citation>Premoli Silva, I., Wade, B. S., and Pearson, P. N.: Taxonomy of
<italic>Globigerinatheka</italic> and <italic>Orbulinoides</italic>, in: Atlas of Eocene Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and
Berggren, W. A., Cushman
Foundation Special Publication, 41, 169–212, Allen Press, Lawrence, KS, USA, 2006.</mixed-citation></ref>
      <ref id="bib1.bib46"><label>46</label><?label 1?><mixed-citation>
Ramadan, A. M., Abd El-Gaied, I. M., Saber, S. G., and Salama, Y. F.:
Foraminiferal biostratigraphy and paleoenvironment evolution recorded in the
Upper Eocene succession in northeastern Desert, Egypt, J.
Sediment. Environ., 6, 485–512, 2021.</mixed-citation></ref>
      <ref id="bib1.bib47"><label>47</label><?label 1?><mixed-citation>Salama, Y., Sayed, M., Saber, S., and Abd El-Gaied, I. M.: Eocene planktonic
foraminifera from the north Eastern Desert, Egypt: Biostratigraphic,
paleoenvironmental and sequence stratigraphy implications, Palaeontol.
Electron., 24, a11, <ext-link xlink:href="https://doi.org/10.26879/1088" ext-link-type="DOI">10.26879/1088</ext-link>, 2021.</mixed-citation></ref>
      <?pagebreak page161?><ref id="bib1.bib48"><label>48</label><?label 1?><mixed-citation>
Seilacher, A.: Begriff und Bedeutung der Fossil-Lagerstätten: Neues
Jahrbuch für Geologie und Paläontologie, Monatshefte, 1, 34–39,
1970.</mixed-citation></ref>
      <ref id="bib1.bib49"><label>49</label><?label 1?><mixed-citation>
Speijer, R. P., Pälike, H., Hooker, J. J., and Ogg, J. G.: The Paleogene
period, in:  Geologic
Time Scale 2020, edited by: Gradstein, F. M., Ogg, J. G., Schmitz, M. D., and Ogg, G. M., 1087–1140, Elsevier,  Amsterdam, the Netherlands, 2020.</mixed-citation></ref>
      <ref id="bib1.bib50"><label>50</label><?label 1?><mixed-citation>Spezzaferri, S., Olsson, R. K., Hemleben, Ch., Wade, B. S., and Coxall, H. K.:
Taxonomy, biostratigraphy, and phylogeny of Oligocene and lower Miocene
<italic>Globoturborotalita</italic>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and Berggren,
W. A., Cushman Foundation of
Foraminiferal Research, Special Publication, No. 46, 231–268, Allen Press, Lawrence, KS, USA, 2018.</mixed-citation></ref>
      <ref id="bib1.bib51"><label>51</label><?label 1?><mixed-citation>
Strougo, A., Faris, M., Haggag, M. A. Y., Abul-Nasr, R. A., and Gingerich, P. D.:
Planktonic Foraminifera and calcareous nannofossil biostratigraphy through
the Middle to Late Eocene transition at Wadi Hitan, Fayum Province, Egypt,
Contributions from the Museum of Paleontology, University of Michigan, 32,
111–138, 2013.</mixed-citation></ref>
      <ref id="bib1.bib52"><label>52</label><?label 1?><mixed-citation>Wade, B. S.: Planktonic foraminiferal biostratigraphy and mechanisms in the
extinction of <italic>Morozovella</italic> in the late Middle Eocene, Mar. Micropaleontol., 51,
23–38, 2004.</mixed-citation></ref>
      <ref id="bib1.bib53"><label>53</label><?label 1?><mixed-citation>Wade, B. S. and Hernitz Kucenjak, M.: Taxonomy, biostratigraphy, and
phylogeny of Oligocene <italic>Acarinina</italic>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber,
B. T., and Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46, 393–402, Allen Press, Lawrence, KS, USA,
2018.</mixed-citation></ref>
      <ref id="bib1.bib54"><label>54</label><?label 1?><mixed-citation>
Wade, B. S. and Pearson, P. N.: Planktonic foraminiferal turnover, diversity
fluctuations and geochemical signals across the Eocene/Oligocene boundary in
Tanzania, Mar. Micropaleontol., 68, 244–255, 2008.</mixed-citation></ref>
      <ref id="bib1.bib55"><label>55</label><?label 1?><mixed-citation>
Wade, B. S. and Olsson, R. K.: Investigation of pre-extinction dwarfing in
Cenozoic planktonic foraminifera, Palaeogeogr. Palaeocl., 284, 39–46, 2009.</mixed-citation></ref>
      <ref id="bib1.bib56"><label>56</label><?label 1?><mixed-citation>Wade, B. S., Al-Sabouni, N., Hemleben, C., and Kroon, D.: Symbiont bleaching
in fossil planktonic foraminifera, Evol. Ecol., 22, 253–265, 2008.
 </mixed-citation></ref><?xmltex \hack{\newpage}?>
      <ref id="bib1.bib57"><label>57</label><?label 1?><mixed-citation>
Wade, B. S., Pearson, P. N., Berggren, W. A., and Pälike, H.: Review and
revision of Cenozoic tropical planktonic foraminiferal biostratigraphy and
calibration to the geomagnetic polarity and astronomical time scale, Earth
Sci. Rev., 104, 111–142, 2011.</mixed-citation></ref>
      <ref id="bib1.bib58"><label>58</label><?label 1?><mixed-citation>Wade, B. S., Premec-Fucek, V., Kamikuri, S., Bartol, M., Luciani, V., and
Pearson, P. N.: Successive extinctions of muricate planktonic foraminifera
(Morozovelloides and Acarinina) mark the base Priabonian, Newsl.
Stratigr., 45, 245–262, <ext-link xlink:href="https://doi.org/10.1127/0078-0421/2012/0023" ext-link-type="DOI">10.1127/0078-0421/2012/0023</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib59"><label>59</label><?label 1?><mixed-citation>
Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and Berggren, W. A.
(Eds.): Atlas of Oligocene Planktonic Foraminifera, Cushman Foundation
Special Publication, No. 46, 528 pp., Allen Press, Lawrence, KS, USA, 2018a.</mixed-citation></ref>
      <ref id="bib1.bib60"><label>60</label><?label 1?><mixed-citation>Wade, B. S., Pearson, P. N., Olsson, R. K., Fraass, A. Leckie, R. M., and
Hemleben, Ch.: Taxonomy, biostratigraphy, and phylogeny of Oligocene and
lower Miocene <italic>Dentoglobigerina</italic> and <italic>Globoquadrina</italic>, in: Atlas of Oligocene Planktonic Foraminifera, edited by:  Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T.,
and Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46,
331–384, 2018b.</mixed-citation></ref>
      <ref id="bib1.bib61"><label>61</label><?label 1?><mixed-citation>Wade, B. S., Olsson, R. K., Pearson, P. N., Edgar, K. M., and Premoli Silva, I.:
Taxonomy, biostratigraphy, and phylogeny of Oligocene <italic>Subbotina</italic>, in: Atlas of Oligocene
Planktonic Foraminifera, edited by: Wade, B. S., Olsson,
R. K., Pearson, P. N., Huber, B. T., and Berggren, W. A., Cushman Foundation of Foraminiferal Research,
Special Publication, No. 46, 307–330, Allen Press, Lawrence, KS, USA, 2018c.</mixed-citation></ref>
      <ref id="bib1.bib62"><label>62</label><?label 1?><mixed-citation>
Wallace, C. A., Dini, S. M., and Al-Farasani, A. N.: Geological map of part of the
Turayf Quadrangle, Sheet 31C, and An Nabk Quadrangle, Sheet 31B, Kingdom of
Saudi Arabia, Ministry of Petroleum and Mineral resources, Saudi Geological
Survey, Geoscience Map Series GM-125C, 1994.</mixed-citation></ref>
      <ref id="bib1.bib63"><label>63</label><?label 1?><mixed-citation>Westerhold, T., Röhl, U., Pälike, H., Wilkens, R., Wilson, P. A., and Acton, G.: Orbitally tuned timescale and astronomical forcing in the middle Eocene to early Oligocene, Clim. Past, 10, 955–973, <ext-link xlink:href="https://doi.org/10.5194/cp-10-955-2014" ext-link-type="DOI">10.5194/cp-10-955-2014</ext-link>, 2014.</mixed-citation></ref>

  </ref-list></back>
    <!--<article-title-html>Upper Eocene planktonic foraminifera from northern Saudi Arabia: implications for stratigraphic ranges</article-title-html>
<abstract-html><p>The Rashrashiyah Formation of the Sirhan Basin in northern Saudi
Arabia contains diverse assemblages of planktonic foraminifera. We examined
the biostratigraphy, stratigraphic range and preservation of upper Eocene
planktonic foraminifera. Assemblages are well-preserved and diverse, with
40 species and 11 genera. All samples are assigned to the Priabonian
<i>Globigerinatheka semiinvoluta</i> Highest Occurrence Zone (E14), consistent with calcareous nannofossil
biostratigraphy indicating Zone CNE17. Well-preserved planktonic
foraminifera assemblages from the lower part of the upper Eocene are rare
worldwide. Our study provides new insights into the stratigraphic ranges of
many species. We find older (Zone E14) stratigraphic occurrences of several
species of <i>Globoturborotalita</i> previously thought to have evolved in the latest Eocene (Zone
E15, E16) or Oligocene; these include <i>G. barbula, G. cancellata, G. gnaucki, G. pseudopraebulloides</i>, and <i>G. paracancellata</i>. Older stratigraphic
occurrences for <i>Dentoglobigerina taci</i> and <i>Subbotina projecta</i> are also found, and <i>Globigerinatheka kugleri</i> occurs at a younger stratigraphic
level than previously proposed. Our revisions to stratigraphic ranges
indicate that the late Eocene had a higher tropical–subtropical diversity of
planktonic foraminifera than hitherto reported.</p></abstract-html>
<ref-html id="bib1.bib1"><label>1</label><mixed-citation>
Agnini, C., Fornaciari, E., Giusberti, L., Grandesso, P., Lanci, L., Luciani, V., Muttoni, G., Palike, H., Rio, D., Spofforth, D. J. A., and Stefani, C.:  Integrated biomagnetostratigraphy of the Alano section (NE Italy): A proposal for defining the middle-late Eocene boundary, Geol. Soc. Am. B., 123, 841–872, 2011.
</mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>2</label><mixed-citation>
Agnini, C., Fornaciari, E., Raffi, I., Catanzariti, R., Pälike, H.,
Backman, J., and Rio, D.: Biozonation and biochronology of Paleogene calcareous
nannofossils from low and middle latitudes, Newsl. Stratigr., 47,
131–181, 2014.
</mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>3</label><mixed-citation>
Agnini, C., Backman, J., Boscolo-Galazzo, F., Condon, D. J., Fornaciari, E.,
Galeotti, S., Giusberti, L., Grandesso, P., Lanci, L., Luciani, V., Monechi,
S., Muttoni, G., Pälike, H., Pampaloni, M. L., Papazzoni, C. A., Pearson,
P. N., Pignatti, J., Premoli Silva, I., Raffi, I., Rio, D., Rook, L., Sahy,
D., Spofforth, D. J. A., Stefani, C., and Wade, B. S.: Proposal for the Global
Boundary Stratotype Section and Point (GSSP) for the Priabonian Stage
(Eocene) at the Alano section (Italy), Episodes, 44, 151–173, 2021.
</mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>4</label><mixed-citation>
Aljahdali, M. H., Elhag, M., Mufrreh, Y., Memesh, A., AlSoubhi, S., and
Zalmout, I. S.: Upper Eocene calcareous nannofossil biostratigraphy: a new
preliminary Priabonian record from northern Saudi Arabia, Appl. Ecol.
Environ. Res., 18, 5607–5625, <a href="https://doi.org/10.15666/aeer/1804_56075625" target="_blank">https://doi.org/10.15666/aeer/1804_56075625</a>, 2020.
</mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>5</label><mixed-citation>
Al-Rawi, M. M.: Petroleum systems in Jordan, GEO ExPro, 11, 1, available at: <a href="https://archives.datapages.com/data/geo-expro-magazine/011/011001/pdfs/22.htm" target="_blank"/> (last access: 26 September 2021), 2014.
</mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>6</label><mixed-citation>
Aze, T., Ezard, T. H. G., Purvis, A., Coxall, H. K., Stewart, D. R. M., Wade, B. S.,
and Pearson, P. N.: A phylogeny of Cenozoic macroperforate planktonic
foraminifera from fossil data, Biol. Rev., 86, 900–927, 2011.
</mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>7</label><mixed-citation>
Berggren, W. A. and Pearson, P. N.: A revised tropical to subtropical planktonic
foraminiferal zonation of the Eocene and Oligocene, J. Foramin.
Res., 35, 279–298, 2005.
</mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>8</label><mixed-citation>
Berggren, W. A., Kent, D. V., Swisher III, C. C., and Aubry, M.-P.: A Revised
Cenozoic Geochronology and Chronostratigraphy, in:
Geochronology, Time Scales and Global Stratigraphic Correlation, edited by: Berggren, W. A., Kent, D. V., Aubry, M.-P., and
Hardenbol, J., SEPM
Society for Sedimentary Geology, Tulsa, OK, USA, 1995.
</mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>9</label><mixed-citation>
Berggren, W. A., Pearson, P. N., Huber, B. T., and Wade, B. S.: Taxonomy,
Biostratigraphy and Phylogeny of Eocene <i>Acarinina</i>,  in: Atlas of Eocene
Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K.,
Huber, B. T., Hemleben, C., and Berggren, W. A., Cushman Foundation Special Publication, 41,
257–326, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>10</label><mixed-citation>
Bown, P. R., Dunkley Jones, T., Lees, J. A., Randell, R. D., Mizzi, J. A.,
Pearson, P. N., Coxall, H. K., Young, J. R., Nicholas, C. J., Karega, A.,
Singano, J., and Wade, B. S.: A Paleogene calcareous microfossil
Konservat-Lagerstätte from the Kilwa Group of coastal Tanzania,
Geol. Soc. Am. Bull., 120, 3–12, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib11"><label>11</label><mixed-citation>
Canudo, J. I. and Molina, E.: Bioestratigrafía con foraminíferos
planctónicos del Paleógeno del Pirineo. in h. Luterbacher (edn.),
Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 186,
97–135, 1992.
</mixed-citation></ref-html>
<ref-html id="bib1.bib12"><label>12</label><mixed-citation>
Cotton, L. J., Zakrevskaya, E. Y., van der Boon, A., Asatryan, G., Hayrapetyan,
F., Israyelyan, A., Krijgsman, W., Less, G., Monechi, S., Papazzoni, C. A.,
Pearson, P. N., Razumovskiy, A., Renema, W., Shcherbinina, E., and Wade,
B. S.: Integrated stratigraphy of the Priabonian (upper Eocene) Urtsadzor
section, Armenia, Newsl. Stratigr., 50, 269–295, <a href="https://doi.org/10.1127/nos/2016/0313" target="_blank">https://doi.org/10.1127/nos/2016/0313</a>, 2017.
</mixed-citation></ref-html>
<ref-html id="bib1.bib13"><label>13</label><mixed-citation>
Dunkley Jones, T., Bown, P. R., Pearson, P. N., Wade, B. S., Coxall, H. K., and
Lear, C. H.: Major shifts in calcareous phytoplankton assemblages through the
Eocene-Oligocene transition of Tanzania and their implications for
low-latitude primary production, Paleoceanography, 23, PA4204,
<a href="https://doi.org/10.1029/2008PA001640" target="_blank">https://doi.org/10.1029/2008PA001640</a>, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib14"><label>14</label><mixed-citation>
Ezard, T. H. G., Aze, T., Pearson, P. N., and Purvis, A.: Interplay between
changing climate and species' ecology drives macroevolutionary dynamics,
Science, 332, 349–351, 2011.
</mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>15</label><mixed-citation>
Farouk, S., Faris, M., Ahmad, F., and Powell, J. H.: New microplanktonic
biostratigraphy and depositional sequences across the Middle-Late Eocene and
Oligocene boundaries in eastern Jordan, GeoArabia – Middle East Petroleum
Geosciences, 20, 145–172, 2015.
</mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>16</label><mixed-citation>
Fayolle, F. and Wade, B. S.: The evolution of Eocene planktonic foraminifera
<i>Dentoglobigerina</i>, J. Syst. Palaeontol., 19, 333–376, 2021.
</mixed-citation></ref-html>
<ref-html id="bib1.bib17"><label>17</label><mixed-citation>
Firth, J. V., Eldrett, J. S., Harding, I. C., Coxall, H. K., and Wade, B. S.:
Integrated biomagnetochronology for the Palaeogene of ODP Hole 647A:
implications for correlating palaeoceanographic events from high to low
latitudes, in: Magnetic Methods and the Timing of Geological Processes, edited by: Jovane, L., Herrero-Bervera, E., Hinnov, L. A., and Housen,
B. A.,
Geological Society, London, Special Publications, 373, 29–78, 2013.
</mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>18</label><mixed-citation>
Fordham, B. G., Aze, T., Haller, C., Zehady, A. K., Pearson, P. N., Ogg, J. G.,
and Wade, B. S.: Future-proofing the Cenozoic macroperforate planktonic
foraminifera phylogeny of Aze &amp; others (2011), PLoS ONE, 13, e0204625,
<a href="https://doi.org/10.1371/journal.pone.0204625" target="_blank">https://doi.org/10.1371/journal.pone.0204625</a>, 2018.
</mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>19</label><mixed-citation>
Fraass, A. J., Kelly, D. C., and Peters, S. E.: Macroevolutionary history of the
planktic foraminifera, Annu. Rev. Earth Planet. Sc., 43,
139–166, <a href="https://doi.org/10.1146/annurev-earth-060614-105059" target="_blank">https://doi.org/10.1146/annurev-earth-060614-105059</a>, 2015.
</mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>20</label><mixed-citation>
Guiraud, R., Issawi, B., and Bosworth, W.: Phanerozoic history of Egypt and
surrounding areas, Peri-Tethys Memoir, 6, 469–509, 2001.
</mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>21</label><mixed-citation>
Haggag, M. A.: <i>Globigerina pseudoampliapertura</i> Zone, a new late Eocene planktonic foraminiferal zone (Fayoum
area, Egypt), Neues Jahrbuch für Geologie und Paläontologie,
Monatshefte, Stuttgart, 295–307, 1990.
</mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>22</label><mixed-citation>
Halawani, M.: Stratigraphic column for the Phanerozoic rocks of Saudi
Arabia. A compilation and synthesis with comments – Saudi Geological
Survey Technical Reports, Kingdom of Saudi Arabia SGS-TR-2001-3 2001,
2001.
</mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>23</label><mixed-citation>
Hemleben, C. and Olsson, R. K.: Wall textures of Eocene planktonic
foraminifera, in: Atlas of Eocene Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and
Berggren, W. A., Cushman
Foundation Special Publication, 41, 47–66, Allen Press, Lawrence, KS, USA, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib24"><label>24</label><mixed-citation>
Huber, B. T., Petrizzo, M. R., Young, J., Falzoni, F., Gilardoni, S., Bown,
P. R., and Wade, B. S.: Pforams@mikrotax: A new online taxonomic database for
planktonic foraminifera, Micropalaeontology, 62, 429–438, 2016.
</mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>25</label><mixed-citation>
Kamikuri, S. and Wade, B. S.: Radiolarian magnetobiochronology and faunal
turnover across the middle/late Eocene boundary at Ocean Drilling Program
Site 1052 in the western North Atlantic Ocean, Mar. Micropaleontol.,
88–89, 41–53, 2012.
</mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>26</label><mixed-citation>
Lear, C., Bailey, T. R., Pearson, P. N., Coxall, H. K., and Rosenthal, Y.:
Cooling and ice growth across the Eocene – Oligocene transition, Geology,
36, 251–254, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib27"><label>27</label><mixed-citation>
Lowery, C. M., Bown, P. R., Fraass, A. J., and Hull, P. M.: Ecological response
of plankton to environmental change: thresholds for extinction, Annu.
Rev. Earth Planet. Sc., 48, 403–429, 2020.
</mixed-citation></ref-html>
<ref-html id="bib1.bib28"><label>28</label><mixed-citation>
Luciani, V., Fornaciari, E., Papazzoni, C. A., Dallanave, E., Giusberti, L.,
Stefani, C., and Amante, E.: Integrated stratigraphy at the
Bartonian–Priabonian transition: correlation between shallow benthic and
calcareous plankton zones (Varignano section, northern Italy), Geol.
Soc. Am. Bull., 132, 495–520, 2020.
</mixed-citation></ref-html>
<ref-html id="bib1.bib29"><label>29</label><mixed-citation>
Martini, E.: Standard Tertiary and Quaternary calcareous nannoplankton
zonation. Proc. II Planktonic Conference, Roma, 1970, Tecnoscienza, Roma,
1971.
</mixed-citation></ref-html>
<ref-html id="bib1.bib30"><label>30</label><mixed-citation>
Meissner, C. R., Griffin Jr., M. B., Riddler, G. P., Marcel Van Eck,
Aspinall, N. C., Farasani, A. M., and Dini, S. M.: Preliminary Geologic Map of
the Thaniyat Turayf Quadrangle, Sheet 29c, Kingdom of Saudi Arabia –
Department of the Interior, US Geological Survey, Washington, DC, 1990.
</mixed-citation></ref-html>
<ref-html id="bib1.bib31"><label>31</label><mixed-citation>
Miller, K. G., Browning, J. V., Aubry, M.-P., Wade, B. S., Katz, M. E., Kulpecz,
A. A., and Wright, J. D.: Eocene-Oligocene global climate and sea-level
changes: St. Stephens Quarry, Alabama, Geol. Soc. Am. Bull.,
120, 34–53, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib32"><label>32</label><mixed-citation>
Moore, T. C. and Kamikuri, S.: Data report: Radiolarian stratigraphy across
the Eocene/Oligocene boundary in the equatorial Pacific, Sites 1218, U1333,
and U1334, In: Proceedings IODP, vol. 320/321, Pälike et
al., Integr. Ocean Drill. Program Management International, Inc., Tokyo,
2012.
</mixed-citation></ref-html>
<ref-html id="bib1.bib33"><label>33</label><mixed-citation>
Norris, R. D., Wilson, P. A., Blum, P., and the Expedition 342 Scientists:
Proc. IODP, 342: College Station, TX (Integrated Ocean Drilling Program), College Station, TX, USA,
2014.
</mixed-citation></ref-html>
<ref-html id="bib1.bib34"><label>34</label><mixed-citation>
Okada, H. and Bukry, D.: Supplementary modification and introduction of code
numbers to the low latitude coccolith biostratigraphic zonation (Bukry,
1973; 1975), Mar. Micropaleontol., 5, 321–325, 1980.
</mixed-citation></ref-html>
<ref-html id="bib1.bib35"><label>35</label><mixed-citation>
Pälike, H., Norris, R. D., Herrle, J. O., Wilson, P. A., Coxall, H. K.,
Lear, C. H., Shackleton, N. J., Tripati, A. K., and Wade, B. S.: The heartbeat of
the Oligocene climate system, Science, 314, 1894–1898, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib36"><label>36</label><mixed-citation>
Pearson, P. N. and Burgess, C. E.: Foraminifer test preservation and
diagenesis: comparison of high latitude Eocene sites, Geol. Soc.
Lond. Spec. Publ., 303, 59–72, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib37"><label>37</label><mixed-citation>
Pearson, P. N. and Kučera, M.: Taxonomy, biostratigraphy, and phylogeny
of Oligocene <i>Turborotalita</i>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and
Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46, 385–392,
2018.
</mixed-citation></ref-html>
<ref-html id="bib1.bib38"><label>38</label><mixed-citation>
Pearson, P. N. and Wade, B. S.: Systematic taxonomy of exceptionally
well-preserved planktonic foraminifera from the Eocene/Oligocene boundary of
Tanzania, Cushman Foundation for Foraminiferal Research Special Publication,
No. 45, 1–85, Allen Press, Lawrence, KS, USA,  2015.
</mixed-citation></ref-html>
<ref-html id="bib1.bib39"><label>39</label><mixed-citation>
Pearson, P. N., Ditchfield, P. W., Singano, J., Harcourt-Brown, K. G.,
Nicholas, C. J., Olsson, R. N., Shackleton, N. J., and Hall, M. A.: Warm
tropical sea surface temperatures in the Late Cretaceous and Eocene epochs,
Nature, 413, 481–487, 2001.
</mixed-citation></ref-html>
<ref-html id="bib1.bib40"><label>40</label><mixed-citation>
Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and Berggren, W. A.:
Atlas of Eocene planktonic foraminifera, Cushman Foundation for
Foraminiferal Research Special Publication, 41, 513 pp., Allen Press, Lawrence, KS, USA, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib41"><label>41</label><mixed-citation>
Pearson, P. N., van Dongen, B. E., Nicholas, C. J., Pancost, R. D., Schouten,
S., Singano, J. M., and Wade, B. S.: Stable warm tropical climate through the
Eocene epoch, Geology, 35, 211–214, 2007.
</mixed-citation></ref-html>
<ref-html id="bib1.bib42"><label>42</label><mixed-citation>
Pearson, P. N., McMillan, I. K., Wade, B. S., Dunkley Jones, T., Coxall, H. K.,
Bown, P. R., and Lear, C. H.: Extinction and environmental change across the
Eocene-Oligocene boundary in Tanzania, Geology, 36, 179–182, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib43"><label>43</label><mixed-citation>
Pearson, P. N., Foster, G. L., and Wade, B. S.: Atmospheric carbon dioxide
through the Eocene-Oligocene climate transition, Nature, 461, 1110–1113,
2009.
</mixed-citation></ref-html>
<ref-html id="bib1.bib44"><label>44</label><mixed-citation>
Postuma, J. A.: Manual of planktonic foraminifera, Elsevier for Shell Group,
The Hague, 406 pp., 1971.
</mixed-citation></ref-html>
<ref-html id="bib1.bib45"><label>45</label><mixed-citation>
Premoli Silva, I., Wade, B. S., and Pearson, P. N.: Taxonomy of
<i>Globigerinatheka</i> and <i>Orbulinoides</i>, in: Atlas of Eocene Planktonic Foraminifera, edited by: Pearson, P. N., Olsson, R. K., Huber, B. T., Hemleben, C., and
Berggren, W. A., Cushman
Foundation Special Publication, 41, 169–212, Allen Press, Lawrence, KS, USA, 2006.
</mixed-citation></ref-html>
<ref-html id="bib1.bib46"><label>46</label><mixed-citation>
Ramadan, A. M., Abd El-Gaied, I. M., Saber, S. G., and Salama, Y. F.:
Foraminiferal biostratigraphy and paleoenvironment evolution recorded in the
Upper Eocene succession in northeastern Desert, Egypt, J.
Sediment. Environ., 6, 485–512, 2021.
</mixed-citation></ref-html>
<ref-html id="bib1.bib47"><label>47</label><mixed-citation>
Salama, Y., Sayed, M., Saber, S., and Abd El-Gaied, I. M.: Eocene planktonic
foraminifera from the north Eastern Desert, Egypt: Biostratigraphic,
paleoenvironmental and sequence stratigraphy implications, Palaeontol.
Electron., 24, a11, <a href="https://doi.org/10.26879/1088" target="_blank">https://doi.org/10.26879/1088</a>, 2021.
</mixed-citation></ref-html>
<ref-html id="bib1.bib48"><label>48</label><mixed-citation>
Seilacher, A.: Begriff und Bedeutung der Fossil-Lagerstätten: Neues
Jahrbuch für Geologie und Paläontologie, Monatshefte, 1, 34–39,
1970.
</mixed-citation></ref-html>
<ref-html id="bib1.bib49"><label>49</label><mixed-citation>
Speijer, R. P., Pälike, H., Hooker, J. J., and Ogg, J. G.: The Paleogene
period, in:  Geologic
Time Scale 2020, edited by: Gradstein, F. M., Ogg, J. G., Schmitz, M. D., and Ogg, G. M., 1087–1140, Elsevier,  Amsterdam, the Netherlands, 2020.
</mixed-citation></ref-html>
<ref-html id="bib1.bib50"><label>50</label><mixed-citation>
Spezzaferri, S., Olsson, R. K., Hemleben, Ch., Wade, B. S., and Coxall, H. K.:
Taxonomy, biostratigraphy, and phylogeny of Oligocene and lower Miocene
<i>Globoturborotalita</i>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and Berggren,
W. A., Cushman Foundation of
Foraminiferal Research, Special Publication, No. 46, 231–268, Allen Press, Lawrence, KS, USA, 2018.
</mixed-citation></ref-html>
<ref-html id="bib1.bib51"><label>51</label><mixed-citation>
Strougo, A., Faris, M., Haggag, M. A. Y., Abul-Nasr, R. A., and Gingerich, P. D.:
Planktonic Foraminifera and calcareous nannofossil biostratigraphy through
the Middle to Late Eocene transition at Wadi Hitan, Fayum Province, Egypt,
Contributions from the Museum of Paleontology, University of Michigan, 32,
111–138, 2013.
</mixed-citation></ref-html>
<ref-html id="bib1.bib52"><label>52</label><mixed-citation>
Wade, B. S.: Planktonic foraminiferal biostratigraphy and mechanisms in the
extinction of <i>Morozovella</i> in the late Middle Eocene, Mar. Micropaleontol., 51,
23–38, 2004.
</mixed-citation></ref-html>
<ref-html id="bib1.bib53"><label>53</label><mixed-citation>
Wade, B. S. and Hernitz Kucenjak, M.: Taxonomy, biostratigraphy, and
phylogeny of Oligocene <i>Acarinina</i>, in: Atlas of Oligocene Planktonic Foraminifera, edited by: Wade, B. S., Olsson, R. K., Pearson, P. N., Huber,
B. T., and Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46, 393–402, Allen Press, Lawrence, KS, USA,
2018.
</mixed-citation></ref-html>
<ref-html id="bib1.bib54"><label>54</label><mixed-citation>
Wade, B. S. and Pearson, P. N.: Planktonic foraminiferal turnover, diversity
fluctuations and geochemical signals across the Eocene/Oligocene boundary in
Tanzania, Mar. Micropaleontol., 68, 244–255, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib55"><label>55</label><mixed-citation>
Wade, B. S. and Olsson, R. K.: Investigation of pre-extinction dwarfing in
Cenozoic planktonic foraminifera, Palaeogeogr. Palaeocl., 284, 39–46, 2009.
</mixed-citation></ref-html>
<ref-html id="bib1.bib56"><label>56</label><mixed-citation>
Wade, B. S., Al-Sabouni, N., Hemleben, C., and Kroon, D.: Symbiont bleaching
in fossil planktonic foraminifera, Evol. Ecol., 22, 253–265, 2008.

</mixed-citation></ref-html>
<ref-html id="bib1.bib57"><label>57</label><mixed-citation>
Wade, B. S., Pearson, P. N., Berggren, W. A., and Pälike, H.: Review and
revision of Cenozoic tropical planktonic foraminiferal biostratigraphy and
calibration to the geomagnetic polarity and astronomical time scale, Earth
Sci. Rev., 104, 111–142, 2011.
</mixed-citation></ref-html>
<ref-html id="bib1.bib58"><label>58</label><mixed-citation>
Wade, B. S., Premec-Fucek, V., Kamikuri, S., Bartol, M., Luciani, V., and
Pearson, P. N.: Successive extinctions of muricate planktonic foraminifera
(Morozovelloides and Acarinina) mark the base Priabonian, Newsl.
Stratigr., 45, 245–262, <a href="https://doi.org/10.1127/0078-0421/2012/0023" target="_blank">https://doi.org/10.1127/0078-0421/2012/0023</a>, 2012.
</mixed-citation></ref-html>
<ref-html id="bib1.bib59"><label>59</label><mixed-citation>
Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T., and Berggren, W. A.
(Eds.): Atlas of Oligocene Planktonic Foraminifera, Cushman Foundation
Special Publication, No. 46, 528 pp., Allen Press, Lawrence, KS, USA, 2018a.
</mixed-citation></ref-html>
<ref-html id="bib1.bib60"><label>60</label><mixed-citation>
Wade, B. S., Pearson, P. N., Olsson, R. K., Fraass, A. Leckie, R. M., and
Hemleben, Ch.: Taxonomy, biostratigraphy, and phylogeny of Oligocene and
lower Miocene <i>Dentoglobigerina</i> and <i>Globoquadrina</i>, in: Atlas of Oligocene Planktonic Foraminifera, edited by:  Wade, B. S., Olsson, R. K., Pearson, P. N., Huber, B. T.,
and Berggren, W. A., Cushman
Foundation of Foraminiferal Research, Special Publication, No. 46,
331–384, 2018b.
</mixed-citation></ref-html>
<ref-html id="bib1.bib61"><label>61</label><mixed-citation>
Wade, B. S., Olsson, R. K., Pearson, P. N., Edgar, K. M., and Premoli Silva, I.:
Taxonomy, biostratigraphy, and phylogeny of Oligocene <i>Subbotina</i>, in: Atlas of Oligocene
Planktonic Foraminifera, edited by: Wade, B. S., Olsson,
R. K., Pearson, P. N., Huber, B. T., and Berggren, W. A., Cushman Foundation of Foraminiferal Research,
Special Publication, No. 46, 307–330, Allen Press, Lawrence, KS, USA, 2018c.
</mixed-citation></ref-html>
<ref-html id="bib1.bib62"><label>62</label><mixed-citation>
Wallace, C. A., Dini, S. M., and Al-Farasani, A. N.: Geological map of part of the
Turayf Quadrangle, Sheet 31C, and An Nabk Quadrangle, Sheet 31B, Kingdom of
Saudi Arabia, Ministry of Petroleum and Mineral resources, Saudi Geological
Survey, Geoscience Map Series GM-125C, 1994.
</mixed-citation></ref-html>
<ref-html id="bib1.bib63"><label>63</label><mixed-citation>
Westerhold, T., Röhl, U., Pälike, H., Wilkens, R., Wilson, P. A., and Acton, G.: Orbitally tuned timescale and astronomical forcing in the middle Eocene to early Oligocene, Clim. Past, 10, 955–973, <a href="https://doi.org/10.5194/cp-10-955-2014" target="_blank">https://doi.org/10.5194/cp-10-955-2014</a>, 2014.
</mixed-citation></ref-html>--></article>
