<document id="7807E9DCF321250AE1C96DE9C70EABFE" ID-DOI="10.26879/1433" ID-ISSN="1094-8074" ID-Zenodo-Dep="16969593" ID-ZooBank="6E7554EF-C09B-4860-AC2A-FA1A6FD53B03" IM.bibliography_requiresApprovalFor="GgImagineBatch" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1756296682779" checkinUser="felipe" docAuthor="Martino, Emanuela Di, Rosso, Antonietta, Taylor, Paul D., Chiu, Ruby W. T., Fujita, Kazuhiko, Kitamura, Akihisa &amp; Yasuhara, Moriaki" docDate="2025" docId="373A87F42D59D967FC0AFAAFDF81FB36" docLanguage="en" docName="PalaeontolElectron.28.1.a7.pdf" docOrigin="Palaeontologia Electronica (a 7) 28 (1)" docSource="https://doi.org/10.26879/1433" docStyle="DocumentStyle:322FC57CC99045555090E8A84CFB029A.4:PalaeontolElectron.2018-.journal_article" docStyleId="322FC57CC99045555090E8A84CFB029A" docStyleName="PalaeontolElectron.2018-.journal_article" docStyleVersion="4" docTitle="Exechonella verrucosa" docType="treatment" docVersion="3" lastPageNumber="45" masterDocId="CB03FF8C2D70D94BFF58FF8EDE68FFD1" masterDocTitle="Unveiling the cheilostome bryozoan fauna of Daidokutsu submarine cave (Okinawa, Japan) over the last 7,000 years" masterLastPageNumber="125" masterPageNumber="1" pageNumber="42" updateTime="1756322269072" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-SA-4.0">
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<mods:title id="C3B8833415EE7E9DDD43ED21EBB630A2">Unveiling the cheilostome bryozoan fauna of Daidokutsu submarine cave (Okinawa, Japan) over the last 7,000 years</mods:title>
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<mods:namePart id="3D3C8E5959343F0881BD94C6071CF861">Martino, Emanuela Di</mods:namePart>
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<paragraph id="BF2C36E22D59D962FC0AFAAFDBFDFAEB" blockId="41.[850,1429,1312,1370]" box="[850,1429,1312,1338]" pageId="41" pageNumber="42">
<heading id="E464818E2D59D962FC0AFAAFDBFDFAEB" box="[850,1429,1312,1338]" centered="true" fontSize="10" level="2" pageId="41" pageNumber="42" reason="7">
<taxonomicName id="78934D612D59D962FC0AFAAFDBFDFAEB" ID-CoL="3DKGG" authority="(Canu and Bassler, 1927)" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" box="[850,1429,1312,1338]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="41" pageNumber="42" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D59D962FC0AFAAFDA08FAEB" box="[850,1120,1313,1338]" italics="true" pageId="41" pageNumber="42">Exechonella verrucosa</emphasis>
(Canu and Bassler, 1927)
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="F78965692D59D962FB64FACFDB09F9BE" pageId="41" pageNumber="42" type="description">
<paragraph id="BF2C36E22D59D962FB64FACFDAC4FA8B" blockId="41.[850,1429,1312,1370]" box="[1084,1196,1345,1370]" pageId="41" pageNumber="42">
<figureCitation id="27A82A672D59D962FB64FACFDAC4FA8B" box="[1084,1196,1345,1370]" captionStart="FIGURE 24" captionStartId="43.[190,281,1736,1759]" captionTargetBox="[188,1441,168,1709]" captionTargetId="figure-21@43.[187,1441,167,1709]" captionTargetPageId="43" captionText="FIGURE 24. Exechonella verrucosa (Canu and Bassler, 1927); PMC EDM-Collection J.H.B.131a. A–D, sample 19074. A, tilted lateral view of colony fragment (400 μm) exposing communication pores. B, frontal view of the same fragment (500 μm). Arrows indicate the nipple-like structure typical of the avicularia of this species. C, close-up of fused foramina with frontal projection and nipple-like structure of an avicularium (100 μm). D, close-up of the primary orifice (150 μm). E–F, sample 19125. E, colony fragment general view (500 μm). F, close-up of a preserved peristome with spiny processes, the lateral one bifurcated (200 μm)." figureDoi="http://doi.org/10.5281/zenodo.16969687" httpUri="https://zenodo.org/record/16969687/files/figure.png" pageId="41" pageNumber="42">Figure 24</figureCitation>
</paragraph>
<paragraph id="BF2C36E22D59D962FC10FAE5DAC7FA75" blockId="41.[840,1441,1387,1912]" pageId="41" pageNumber="42">
v. 1927 
<taxonomicName id="78934D612D59D962FC9DFAE5DBFFFA55" authority="Canu and Bassler" authorityName="Canu and Bassler" box="[965,1431,1387,1413]" class="Gymnolaemata" family="Teuchoporidae" genus="Coleopora" kingdom="Animalia" order="Cheilostomatida" pageId="41" pageNumber="42" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D59D962FC9DFAE5DAD3FA55" box="[965,1211,1387,1412]" italics="true" pageId="41" pageNumber="42">Coleopora verrucosa</emphasis>
Canu and Bassler
</taxonomicName>
, p. 6, 42, pl. 1, fig. 7.
</paragraph>
<paragraph id="BF2C36E22D59D962FC10FA3FDAF7F9DB" blockId="41.[840,1441,1387,1912]" pageId="41" pageNumber="42">
v. 1929 
<taxonomicName id="78934D612D59D962FC9DFA3EDBFFFA1B" authority="Canu and Bassler" authorityName="Canu and Bassler" box="[965,1431,1456,1482]" class="Gymnolaemata" family="Teuchoporidae" genus="Coleopora" kingdom="Animalia" order="Cheilostomatida" pageId="41" pageNumber="42" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D59D962FC9DFA3EDAD3FA1B" box="[965,1211,1456,1482]" italics="true" pageId="41" pageNumber="42">Coleopora verrucosa</emphasis>
Canu and Bassler
</taxonomicName>
; Canu and Bassler, p. 267–268, pl. 20, fig. 4, pl. 26, fig. 9.
</paragraph>
<paragraph id="BF2C36E22D59D962FC10F998DB09F9BE" blockId="41.[840,1441,1387,1912]" pageId="41" pageNumber="42">
v. 2017 
<taxonomicName id="78934D612D59D962FC9DF998DA43F99E" authority="(Canu and Bassler)" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="41" pageNumber="42" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D59D962FC9DF998DABAF9FE" box="[965,1234,1558,1583]" italics="true" pageId="41" pageNumber="42">Exechonella verrucosa</emphasis>
(Canu and Bassler)
</taxonomicName>
; Cáceres-Chamizo, Sanner, Tilbrook and Ostrovsky, p. 56, fig. 23.
</paragraph>
</subSubSection>
<subSubSection id="F78965692D59D962FC10F9F5DB74F925" pageId="41" pageNumber="42" type="materials_examined">
<paragraph id="BF2C36E22D59D962FC10F9F5DBC8F944" blockId="41.[840,1441,1387,1912]" box="[840,1440,1659,1685]" pageId="41" pageNumber="42">
<emphasis id="8DE7EAF02D59D962FC10F9F5DA21F945" bold="true" box="[840,1097,1659,1684]" pageId="41" pageNumber="42">Figured material.</emphasis>
PMC EDM-Collection
</paragraph>
<paragraph id="BF2C36E22D59D962FC10F915DB74F925" blockId="41.[840,1441,1387,1912]" pageId="41" pageNumber="42">
J.H.B.131a, sample 19074 (
<figureCitation id="27A82A672D59D962FBF3F915DB34F965" box="[1195,1372,1691,1716]" captionStart="FIGURE 24" captionStartId="43.[190,281,1736,1759]" captionTargetBox="[188,1441,168,1709]" captionTargetId="figure-21@43.[187,1441,167,1709]" captionTargetPageId="43" captionText="FIGURE 24. Exechonella verrucosa (Canu and Bassler, 1927); PMC EDM-Collection J.H.B.131a. A–D, sample 19074. A, tilted lateral view of colony fragment (400 μm) exposing communication pores. B, frontal view of the same fragment (500 μm). Arrows indicate the nipple-like structure typical of the avicularia of this species. C, close-up of fused foramina with frontal projection and nipple-like structure of an avicularium (100 μm). D, close-up of the primary orifice (150 μm). E–F, sample 19125. E, colony fragment general view (500 μm). F, close-up of a preserved peristome with spiny processes, the lateral one bifurcated (200 μm)." figureDoi="http://doi.org/10.5281/zenodo.16969687" httpUri="https://zenodo.org/record/16969687/files/figure.png" pageId="41" pageNumber="42">Figure 24A–D</figureCitation>
) and sample 19125 (
<figureCitation id="27A82A672D59D962FB53F935DAD4F905" box="[1035,1212,1723,1748]" captionStart="FIGURE 24" captionStartId="43.[190,281,1736,1759]" captionTargetBox="[188,1441,168,1709]" captionTargetId="figure-21@43.[187,1441,167,1709]" captionTargetPageId="43" captionText="FIGURE 24. Exechonella verrucosa (Canu and Bassler, 1927); PMC EDM-Collection J.H.B.131a. A–D, sample 19074. A, tilted lateral view of colony fragment (400 μm) exposing communication pores. B, frontal view of the same fragment (500 μm). Arrows indicate the nipple-like structure typical of the avicularia of this species. C, close-up of fused foramina with frontal projection and nipple-like structure of an avicularium (100 μm). D, close-up of the primary orifice (150 μm). E–F, sample 19125. E, colony fragment general view (500 μm). F, close-up of a preserved peristome with spiny processes, the lateral one bifurcated (200 μm)." figureDoi="http://doi.org/10.5281/zenodo.16969687" httpUri="https://zenodo.org/record/16969687/files/figure.png" pageId="41" pageNumber="42">Figure 24E–F</figureCitation>
); Core 19, Daidokutsu cave, 
<collectingRegion id="7D57F8002D59D962FC8EF955DA2AF924" box="[982,1090,1755,1781]" country="Japan" name="Okinawa" pageId="41" pageNumber="42">Okinawa</collectingRegion>
, 
<collectingCountry id="C78476722D59D962FB15F955DAF2F925" box="[1101,1178,1755,1780]" name="Japan" pageId="41" pageNumber="42">Japan</collectingCountry>
, Holocene.
</paragraph>
</subSubSection>
<subSubSection id="F78965692D59D967FC10F971DFB4FD92" lastPageId="44" lastPageNumber="45" pageId="41" pageNumber="42" type="description">
<paragraph id="BF2C36E22D59D962FC10F971DBC9F889" blockId="41.[840,1441,1387,1912]" pageId="41" pageNumber="42">
<emphasis id="8DE7EAF02D59D962FC10F971DD8AF8C9" bold="true" box="[840,994,1791,1816]" pageId="41" pageNumber="42">Description.</emphasis>
Colony encrusting. Autozooids distinct, separated by deep grooves, oval to rhomboidal, slightly longer than wide (mean ZL/ZW 1.13).
</paragraph>
<paragraph id="BF2C36E22D59D967FC10F8D1DC28FE6E" blockId="41.[840,1441,1387,1912]" lastBlockId="44.[192,793,166,1255]" lastPageId="44" lastPageNumber="45" pageId="41" pageNumber="42">
Frontal shield slightly convex, wrinkled, tessellated, evenly and densely punctured by over 40 minute, circular foramina, 
<emphasis id="8DE7EAF02D5AD961FEC4F9DDDFD8F9BD" box="[412,432,1619,1644]" italics="true" pageId="42" pageNumber="43">c.</emphasis>
15 µm in diameter; foramina centrally located within a larger funnel-shaped gymnocystal rim, measuring between 30–40 µm in diameter; often, rims of 2–3 foramina fusing together, forming distinctive reniform to trifoliate clusters adorned with tubular, hollow, spike-like frontal projections, 75–125 µm long, 25–40 µm in diameter at the tip; projections randomly distributed across the frontal shield, with about 6–8 per zooid in the specimens examined; numerous circular marginal pores, roughly 15 µm in diameter, distributed around zooidal periphery. Primary orifice subcircular, slightly wider than long, surrounded by a short, collar-like peristome; stout, triangular condyles, 35–40 µm long by 40–45 µm wide, medially directed, placed at two-thirds of orifice length, separating the orifice into equally wide anter and poster, with anter occupying two-thirds and poster one-third of orifice length. Peristome externally pustulose, internally smooth to slightly wrinkled, forming a sloping shelf proximally, developing spiny processes around its rim; eight such processes counted in the single zooid in which they were all preserved, with the lateral one showing a bifurcated structure. Avicularia present on the rims of lateral or proximal foramina producing tubular projections, characterized by a distinctive nipple-like structure with a central pore.
</paragraph>
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<paragraph id="BF2C36E22D5AD961FF9DFAC2DDA6FA27" blockId="42.[197,1436,1356,1526]" pageId="42" pageNumber="43">
<emphasis id="8DE7EAF02D5AD961FF9DFAC2DF21FAB2" bold="true" box="[197,329,1356,1379]" pageId="42" pageNumber="43">FIGURE 23.</emphasis>
<taxonomicName id="78934D612D5AD961FE08FAC2DC7AFAB2" authorityName="Martino &amp; Rosso &amp; Taylor &amp; Chiu &amp; Fujita &amp; Kitamura &amp; Yasuhara" authorityYear="2025" box="[336,530,1356,1379]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="42" pageNumber="43" phylum="Bryozoa" rank="species" species="juliae" status="sp. nov.">
<emphasis id="8DE7EAF02D5AD961FE08FAC2DC7AFAB2" box="[336,530,1356,1379]" italics="true" pageId="42" pageNumber="43">Exechonella juliae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="96D4578B2D5AD961FD41FAC2DC05FAB2" box="[537,621,1356,1379]" pageId="42" pageNumber="43" rank="species">sp. nov.</taxonomicNameLabel>
Di Martino, Rosso and Taylor. A–B, paratype PMC. B54. 29.7.2024b1, sample 19059. A, colony fragment general view (1 mm). B, close-up of autozooid (500 μm). C, paratype PMC. B54. 29.7.2024b2, sample 19015, close-up of an autozooid with a frontal projection (500 μm). D–E, holotype PMC. B54. 29.7.2024a, sample 19014. D, close-up of an autozooid with frontal projection and view of peristome internal surface (200 μm). E, close-up of the primary orifice (100 μm); F, paratype PMC. B54. 29.7.2024b3, sample 19042, group of zooids showing the flared, jagged rim of the tubular peristomes (500 μm).
</paragraph>
</caption>
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<paragraph id="BF2C36E22D5BD960FFE6F946DCC7F8A3" blockId="43.[190,1439,1736,1906]" pageId="43" pageNumber="44">
<emphasis id="8DE7EAF02D5BD960FFE6F946DF28F90E" bold="true" box="[190,320,1736,1759]" pageId="43" pageNumber="44">FIGURE 24.</emphasis>
<taxonomicName id="78934D612D5BD960FE1EF946DD38F90E" authority="(Canu and Bassler, 1927)" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" box="[326,848,1736,1759]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="43" pageNumber="44" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D5BD960FE1EF946DC50F90E" box="[326,568,1736,1759]" italics="true" pageId="43" pageNumber="44">Exechonella verrucosa</emphasis>
(Canu and Bassler, 1927)
</taxonomicName>
; PMC EDM-Collection J.H.B.131a. A–D, sample 19074. A, tilted lateral view of colony fragment (400 μm) exposing communication pores. B, frontal view of the same fragment (500 μm). Arrows indicate the nipple-like structure typical of the avicularia of this species. C, close-up of fused foramina with frontal projection and nipple-like structure of an avicularium (100 μm). D, close-up of the primary orifice (150 μm). E–F, sample 19125. E, colony fragment general view (500 μm). F, close-up of a preserved peristome with spiny processes, the lateral one bifurcated (200 μm).
</paragraph>
</caption>
<paragraph id="BF2C36E22D5CD967FF98FE44DFB4FD92" blockId="44.[192,793,166,1255]" pageId="44" pageNumber="45">
<emphasis id="8DE7EAF02D5CD967FF98FE44DFABFE32" bold="true" box="[192,451,458,483]" pageId="44" pageNumber="45">Measurements (µm).</emphasis>
ZL 1159±96, 1059–1251 (2, 3); ZW 1028±59, 975–1092 (2, 3); OL 275±3, 273– 278 (2, 3); OW 322±26, 298–349 (2, 3); PeL 221±55, 182–260 (2, 2).
</paragraph>
</subSubSection>
<subSubSection id="F78965692D5CD967FF98FDC0DF81FB36" pageId="44" pageNumber="45" type="discussion">
<paragraph id="BF2C36E22D5CD967FF98FDC0DF81FB36" blockId="44.[192,793,166,1255]" pageId="44" pageNumber="45">
<emphasis id="8DE7EAF02D5CD967FF98FDC0DF50FDB6" bold="true" box="[192,312,590,615]" pageId="44" pageNumber="45">Remarks.</emphasis>
To date, 
<taxonomicName id="78934D612D5CD967FE8AFDC0DC87FDB6" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" box="[466,751,590,615]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="44" pageNumber="45" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D5CD967FE8AFDC0DC87FDB6" box="[466,751,590,615]" italics="true" pageId="44" pageNumber="45">Exechonella verrucosa</emphasis>
</taxonomicName>
is unequivocally known only from the 
<collectingCountry id="C78476722D5CD967FDD7FDE0DD7CFD56" box="[655,788,622,647]" name="Philippines" pageId="44" pageNumber="45">Philippines</collectingCountry>
. Records from other geographical areas have been reassigned to new species in an effort to clarify the 
<taxonomicName id="78934D612D5CD967FF98FD40DF33FD36" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" box="[192,347,718,743]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="44" pageNumber="45" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D5CD967FF98FD40DF33FD36" box="[192,347,718,743]" italics="true" pageId="44" pageNumber="45">E. verrucosa</emphasis>
</taxonomicName>
species-complex (Cáceres-Chamizo et al., 2017). Due to the limited availability of material, consisting of only a few fragments with one or two zooids each, it is challenging to observe morphological variation. However, the key characteristics of our specimens fit the updated description of 
<taxonomicName id="78934D612D5CD967FF98FC00DF37FC76" baseAuthorityName="Canu and Bassler" baseAuthorityYear="1927" box="[192,351,910,935]" class="Gymnolaemata" family="Exechonellidae" genus="Exechonella" kingdom="Animalia" order="Cheilostomatida" pageId="44" pageNumber="45" phylum="Bryozoa" rank="species" species="verrucosa">
<emphasis id="8DE7EAF02D5CD967FF98FC00DF37FC76" box="[192,351,910,935]" italics="true" pageId="44" pageNumber="45">E. verrucosa</emphasis>
</taxonomicName>
, which appears to be the closest match among those described within this genus, also in terms of size, number and distribution of foramina. Notable differences we have observed compared to the Recent 
<typeStatus id="602888402D5CD967FEA1FB80DC43FBF6" box="[505,555,1038,1063]" pageId="44" pageNumber="45">type</typeStatus>
specimens include the absence of well-defined longitudinal striations on the inner surface of the peristome; instead, only wrinkles concentric to the proximal orifice margin have been observed. Additionally, adventitious kenozooids associated with the avicularia are also absent in our specimens.
</paragraph>
</subSubSection>
</treatment>
</document>