treatments-xml/data/6F/0C/24/6F0C243FFF80256FFCD8667F0336FEFE.xml
2024-06-21 12:39:53 +02:00

259 lines
52 KiB
XML
Raw Blame History

This file contains ambiguous Unicode characters

This file contains Unicode characters that might be confused with other characters. If you think that this is intentional, you can safely ignore this warning. Use the Escape button to reveal them.

<document id="F0B8D90B0A9FCD9E62644CD0403A0B24" ID-DOI="10.1111/j.1096-3642.2005.00179.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5434718" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630433379954" checkinUser="diego" docAuthor="Ostrovsky, Andrew N. &amp; Taylor, Paul D." docDate="2005" docId="6F0C243FFF80256FFCD8667F0336FEFE" docLanguage="en" docName="j.1096-3642.2005.00179.x.pdf" docOrigin="Zoological Journal of the Linnean Society 144 (3)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2005.00179.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Tendra zostericola Nordmann, NHM 1839" docType="treatment" docVersion="5" lastPageNumber="322" masterDocId="93355C47FF82256AFFA867060711FFE6" masterDocTitle="Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans" masterLastPageNumber="361" masterPageNumber="317" pageNumber="319" updateTime="1699269389577" updateUser="plazi" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
<mods:mods id="85AB7594BFA1A94C17B089CBC70FCD95" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="D0ACAA963F2A9DFFBB05B1E7F0BA760B">
<mods:title id="7E6E548C8B6A91887C1F82A223A8CF48">Brood chambers constructed from spines in fossil and Recent cheilostome bryozoans</mods:title>
</mods:titleInfo>
<mods:name id="4D663DBCD3293F3B4DFE6D12E195EE3E" type="personal">
<mods:role id="6F880E8082E8BC78D788511A87DB7FAB">
<mods:roleTerm id="1BB04C6E7A31E38753589BEBF7F5FF30">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="8F0AB11FA50C3B28A2060F586EE22550">Ostrovsky, Andrew N.</mods:namePart>
</mods:name>
<mods:name id="F42A4839DA21251B5EEBC12FA3188F28" type="personal">
<mods:role id="DBD35D979E5678EA216BCF16C4FB71EB">
<mods:roleTerm id="B1C4C5A2F70633C277389898CA03A364">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="DDF47340C41E91288E1F0E737F106010">Taylor, Paul D.</mods:namePart>
</mods:name>
<mods:typeOfResource id="11129FE233AAB3C6F655CB4C3A8CFE94">text</mods:typeOfResource>
<mods:relatedItem id="913E03A2C0AF00AF6054374FAB632F9E" type="host">
<mods:titleInfo id="3A4ABC71D76672CEBBB9ED6E2DE98079">
<mods:title id="4AF48CCAA9177764A6B1F641443D95DC">Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part id="06EC0B69F23F57B9AFF231B74BBFFBC3">
<mods:date id="AB9D36F775C3CF7A874D761E4DA37653">2005</mods:date>
<mods:detail id="3400A1B7B8413261DB3D802CD4BC7720" type="pubDate">
<mods:number id="EDFB129516DF6F114C21CD16070BB951">2005-07-31</mods:number>
</mods:detail>
<mods:detail id="F2070A28AD1D505D8FFBDCE0DDAB0E7A" type="volume">
<mods:number id="242B13939D7A994F9BD028633A1C1528">144</mods:number>
</mods:detail>
<mods:detail id="83842A443911FBB06A184BCEC417E973" type="issue">
<mods:number id="33E0C709326773EABFD9838BEF66DF1E">3</mods:number>
</mods:detail>
<mods:extent id="DB3157FE800F487124499ECB4567FC0A" unit="page">
<mods:start id="A2CD0034D3EDC629192CF83A122F3911">317</mods:start>
<mods:end id="6AD47D39942AD65F8434DA83E189867F">361</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location id="8426F9C11332B6562423F3AFCC3330D0">
<mods:url id="93B966235C456B75DE7B1F917219895F">https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2005.00179.x</mods:url>
</mods:location>
<mods:classification id="1852A8744A720AA6ACE2DEB8A274A627">journal article</mods:classification>
<mods:identifier id="69D3975FC80B8C66F3AC44A81F0CA2D0" type="DOI">10.1111/j.1096-3642.2005.00179.x</mods:identifier>
<mods:identifier id="EFB93C1C04EFB2951513054A9C050C9E" type="ISSN">0024-4082</mods:identifier>
<mods:identifier id="BA7649092A24781527158ED419E80A05" type="Zenodo-Dep">5434718</mods:identifier>
</mods:mods>
<treatment id="6F0C243FFF80256FFCD8667F0336FEFE" LSID="urn:lsid:plazi:treatment:6F0C243FFF80256FFCD8667F0336FEFE" httpUri="http://treatment.plazi.org/id/6F0C243FFF80256FFCD8667F0336FEFE" lastPageId="5" lastPageNumber="322" pageId="2" pageNumber="319">
<subSubSection id="AFBFC6A2FF802568FCD8667F0252FE76" box="[880,1347,376,400]" pageId="2" pageNumber="319" type="nomenclature">
<paragraph id="E71A9529FF802568FCD8667F0252FE76" blockId="2.[880,1347,312,432]" box="[880,1347,376,400]" pageId="2" pageNumber="319">
<heading id="BC522245FF802568FCD8667F0252FE76" box="[880,1347,376,400]" centered="true" fontSize="9" level="2" pageId="2" pageNumber="319" reason="2">
<taxonomicName id="20A5EEAAFF802568FCD8667F0252FE76" ID-CoL="7BQV6" authority="NORDMANN, 1839" authorityName="Nordmann, NHM" authorityYear="1839" box="[880,1347,376,400]" class="Gymnolaemata" family="Tendridae" genus="Tendra" kingdom="Animalia" order="Cheilostomatida" pageId="2" pageNumber="319" phylum="Bryozoa" rank="species" species="zostericola">
<emphasis id="D5D1493BFF802568FCD8667F037EFE69" box="[880,1135,377,400]" italics="true" pageId="2" pageNumber="319">TENDRA ZOSTERICOLA</emphasis>
<bibRefCitation id="8334E8D8FF802568FBDF667E0252FE76" author="Nordmann MA" box="[1143,1347,376,400]" pageId="2" pageNumber="319" pagination="185 - 191" refId="ref27386" refString="Nordmann MA. 1839. Recherches microscopiques sur l'anatomie et le developpement du Tendra zostericola, espece de polype de la section des Bryozoaires. Annales des Sciences Naturelles, Serie 2 (11): 185 - 191." type="journal article" year="1839">NORDMANN, 1839</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="AFBFC6A2FF80256FFC5B669E0336FEFE" lastPageId="5" lastPageNumber="322" pageId="2" pageNumber="319" type="description">
<paragraph id="E71A9529FF802568FC5B669E03D0FE56" blockId="2.[880,1347,312,432]" box="[1011,1217,408,432]" pageId="2" pageNumber="319">
(
<figureCitation id="7F9E89ACFF802568FC53669E036CFE56" box="[1019,1149,408,432]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">FIGS 1A- D</figureCitation>
,
<figureCitation id="7F9E89ACFF802568FB22669E03ABFE56" box="[1162,1210,408,432]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="2" pageNumber="319">21A</figureCitation>
)
</paragraph>
<paragraph id="E71A9529FF802568FC8E66CF03CCFE18" blockId="2.[806,1422,457,510]" pageId="2" pageNumber="319">
<emphasis id="D5D1493BFF802568FC8E66CF049EFE38" box="[806,911,457,478]" italics="true" pageId="2" pageNumber="319">Material:</emphasis>
NHM 11.10.1.489, Recent, Sebastopol Bay, Black Sea, collected by A. Ostroumoff.
</paragraph>
<paragraph id="E71A9529FF802568FC8E651004F8FD05" blockId="2.[806,1423,534,1905]" pageId="2" pageNumber="319">
<emphasis id="D5D1493BFF802568FC8E651004BFFDCD" box="[806,942,534,555]" italics="true" pageId="2" pageNumber="319">Description:</emphasis>
Non-brooding autozooids possess two oral and sometimes one or two, occasionally more, proximal mural spines that are more slender and shorter than the oral spines (
<figureCitation id="7F9E89ACFF802568FBD5657403E5FD61" box="[1149,1268,626,648]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1A, D</figureCitation>
). Oral spines are truncated, their tips probably having a terminal membranous window (cf.
<bibRefCitation id="8334E8D8FF802568FBC065A9023BFD23" author="Ostrovsky AN" box="[1128,1322,687,709]" pageId="2" pageNumber="319" pagination="287 - 318" refId="ref27546" refString="Ostrovsky AN. 1998. Comparative studies of ovicell anatomy and reproductive patterns in Cribrilina annulata and Celleporella hyalina (Bryozoa: Cheilostomatida). Acta Zoologica 79 (4): 287 - 318." type="journal article" year="1998">Ostrovsky, 1998</bibRefCitation>
). Mural spines are acute.
</paragraph>
<paragraph id="E71A9529FF802568FC9765EB047CFAEA" blockId="2.[806,1423,534,1905]" pageId="2" pageNumber="319">
In addition to the oral spines, the left and right sides of the mural rim in brooding autozooids have a row of horizontally inclined, basally flattened spines (
<figureCitation id="7F9E89ACFF802568FC86644E04D2FCB8" box="[814,963,840,862]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1A, B, D</figureCitation>
). These are long and acute, overarching the frontal membrane to give a brooding cavity between this membrane and the undersides of the spines (
<figureCitation id="7F9E89ACFF802568FC2964A204F3FC5C" box="[897,994,932,954]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="2" pageNumber="319">Fig. 21A</figureCitation>
). Each row consists of 1015 spines (up to 17, according to
<bibRefCitation id="8334E8D8FF802568FB8264C503F7FC3F" author="Repiachoff W" box="[1066,1254,963,985]" pageId="2" pageNumber="319" pagination="129 - 142" refId="ref27910" refString="Repiachoff W. 1875. Zur Entwickelungsgeschichte der Tendra zostericola. Zeitschrift fur Wissenschaftliche Zoologie 25: 129 - 142." type="journal article" year="1875">Repiachoff, 1875</bibRefCitation>
and
<bibRefCitation id="8334E8D8FF802568FAB764C5044EFC11" author="Levinsen GMR" pageId="2" pageNumber="319" refId="ref27042" refString="Levinsen GMR. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa. Copenhagen: Bagge F." type="book" year="1909">Levinsen, 1909</bibRefCitation>
; 1318, according to
<bibRefCitation id="8334E8D8FF802568FBD464E404D6FBF0" author="Occhipinti Ambrogi A &amp; d'Hondt J-L" pageId="2" pageNumber="319" pagination="191 - 198" refId="ref27426" refString="Occhipinti Ambrogi A, d'Hondt J-L. 1981. Distribution of bryozoans in brackish waters of Italy. In: Larwood GP, Nielsen C, eds. Recent and fossil Bryozoa. Fredensborg: Olsen &amp; Olsen, 191 - 198." type="book chapter" year="1981">Occhipinti Ambrogi &amp; dHondt, 1981</bibRefCitation>
), growing towards the opposite row. The proximal part of the opesia is devoid of spines, leaving a gap for oviposition of the eggs and for release of the larvae. This entrance to the brooding cavity may be open but in most instances it is plugged by the operculum of the maternal zooid (
<figureCitation id="7F9E89ACFF802568FBDE639C03D8FB56" box="[1142,1225,1178,1200]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1A</figureCitation>
), similarly to the so-called cleithral ovicells found in neocheilostomes. The brood chamber has two openings, proximal and distal.
</paragraph>
<paragraph id="E71A9529FF802569FC97621205E3FA25" blockId="2.[806,1423,534,1905]" lastBlockId="3.[162,778,1392,1905]" lastPageId="3" lastPageNumber="320" pageId="2" pageNumber="319">
During development of the brood chamber, the proximal spines evidently commence growth before the distal spines (
<figureCitation id="7F9E89ACFF802568FC6F6254032AFA8E" box="[967,1083,1362,1384]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1A, C</figureCitation>
). They meet the spines growing towards them from the opposite side along the midline of the zooidal frontal surface. In some instances spine growth immediately ceases at this point, but in other cases the spines overgrow one another, sometimes even reaching the opposite side of the mural rim (
<figureCitation id="7F9E89ACFF802568FC4E610C0329F9C6" box="[998,1080,1546,1568]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1B</figureCitation>
). Spine arrangement varies, from rather regular to chaotic, and from very tight (though not fused laterally) to loose with slits between adjacent spines (
<figureCitation id="7F9E89ACFF802568FC516160035FF99A" box="[1017,1102,1638,1660]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1D</figureCitation>
). Within the same brood chamber, spines can also be very variable in shape: wide or narrow, flat or more cylindrical, unbranched or bifid (
<figureCitation id="7F9E89ACFF802568FCC261C404ADF93E" box="[874,956,1730,1752]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1C</figureCitation>
), straight or curved, and growing along or above the spine lattice. In rare examples only one spine row is developed, either on the left or right side of the zooid (
<figureCitation id="7F9E89ACFF802568FC1D60180319F8D2" box="[949,1032,1822,1844]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="2" pageNumber="319">Fig. 1C</figureCitation>
). In addition, some zooids have up to nine thin mural spines of different length which are not in contact with the spines from either the same or the opposite side of the zooid (
<figureCitation id="7F9E89ACFF812569FD9B62760597FA60" box="[563,646,1392,1414]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="3" pageNumber="320">Fig. 1D</figureCitation>
) (see also
<bibRefCitation id="8334E8D8FF812569FF0A62890640FA43" author="Levinsen GMR" box="[162,337,1423,1445]" pageId="3" pageNumber="320" refId="ref27042" refString="Levinsen GMR. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa. Copenhagen: Bagge F." type="book" year="1909">Levinsen, 1909</bibRefCitation>
). It is difficult to envisage such a loose construction serving as an effective brood chamber.
</paragraph>
<caption id="B3DAC5A1FF812569FF0A63990630FACE" ID-DOI="http://doi.org/10.5281/zenodo.5434720" ID-Zenodo-Dep="5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="3" pageNumber="320" startId="3.[162,240,1183,1202]" targetBox="[162,1431,198,1150]" targetPageId="3">
<paragraph id="E71A9529FF812569FF0A63990630FACE" blockId="3.[162,1442,1183,1320]" pageId="3" pageNumber="320">
<emphasis id="D5D1493BFF812569FF0A6399061BFB54" bold="true" box="[162,266,1183,1202]" pageId="3" pageNumber="320">Figure 1.</emphasis>
<taxonomicName id="20A5EEAAFF812569FEBE63990585FB54" authority="Nordmann, NHM" authorityName="Nordmann, NHM" authorityYear="1839" box="[278,660,1183,1202]" class="Gymnolaemata" family="Tendridae" genus="Tendra" kingdom="Animalia" order="Cheilostomatida" pageId="3" pageNumber="320" phylum="Bryozoa" rank="species" species="zostericola">
<emphasis id="D5D1493BFF812569FEBE639906C5FB54" box="[278,468,1183,1202]" italics="true" pageId="3" pageNumber="320">Tendra zostericola</emphasis>
Nordmann, NHM
</taxonomicName>
11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm.
</paragraph>
</caption>
<paragraph id="E71A9529FF812569FF1362CA06F9F931" blockId="3.[162,778,1392,1905]" pageId="3" pageNumber="320">
All of the oral and mural spines are articulated, as indicated by the presence of a ring furrow at the base of each spine. In contrast, brood-chamber spines appear not to be articulated; unlike oral and mural spines, none are broken off basally. However, sections of decalcified material are required to confirm this supposition. The cavities of all spine
<typeStatus id="381E2B8BFF812569FDE56183059BF97C" box="[589,650,1669,1690]" pageId="3" pageNumber="320">types</typeStatus>
are apparently confluent with the visceral coelom of the zooid distal of the maternal zooid.
</paragraph>
<paragraph id="E71A9529FF812569FF1361E6061DF897" blockId="3.[162,778,1392,1905]" pageId="3" pageNumber="320">
Brood chambers are formed near the distal margins of colonies, often being located at the growing edge. In many instances several brooding zooids are found in the same longitudinal row, one following the other (
<figureCitation id="7F9E89ACFF812569FF02605D07EFF897" box="[170,254,1883,1905]" captionStart="Figure 1" captionStartId="3.[162,240,1183,1202]" captionTargetBox="[162,1431,198,1150]" captionTargetId="figure-389@3.[162,1432,197,1150]" captionTargetPageId="3" captionText="Figure 1. Tendra zostericola Nordmann, NHM 11.10.1.489, Recent, Black Sea. A, part of a colony with both brooding and nonbrooding zooids. B, brooding zooid with overlapping frontal spines. C, partially formed brood chamber (left) and brooding zooid with spines developed only on one side (right). D, sparse mural spines in nonbrooding zooid (lower right) and three brooding zooids with different variants of the frontal spine arrangement. Scale bars: A = 250 Mm; B = 100 Mm; C = 125 Mm; D = 150 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434720" httpUri="https://zenodo.org/record/5434720/files/figure.png" pageId="3" pageNumber="320">Fig. 1D</figureCitation>
).
</paragraph>
<paragraph id="E71A9529FF812569FC92627604B2F8D5" blockId="3.[826,1442,1392,1905]" pageId="3" pageNumber="320">
<emphasis id="D5D1493BFF812569FC92627604B7FA63" box="[826,934,1392,1413]" italics="true" pageId="3" pageNumber="320">Remarks:</emphasis>
The term acanthostegal or acanthostegous ooecia was introduced by
<bibRefCitation id="8334E8D8FF812569FBDC62890239FA42" author="Levinsen GMR" box="[1140,1320,1423,1445]" pageId="3" pageNumber="320" pagination="1 - 31" refId="ref27019" refString="Levinsen GMR. 1902. Studies on Bryozoa. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 54: 1 - 31." type="journal article" year="1902">Levinsen (1902</bibRefCitation>
,
<bibRefCitation id="8334E8D8FF812569FA9062890261FA43" author="Levinsen GMR" box="[1336,1392,1423,1445]" pageId="3" pageNumber="320" refId="ref27042" refString="Levinsen GMR. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa. Copenhagen: Bagge F." type="book" year="1909">1909</bibRefCitation>
) for the unusual brood chambers constructed of spines in
<taxonomicName id="20A5EEAAFF812569FC9262CA030DFA07" authorityName="Nordmann, NHM" authorityYear="1839" box="[826,1052,1484,1505]" class="Gymnolaemata" family="Tendridae" genus="Tendra" kingdom="Animalia" order="Cheilostomatida" pageId="3" pageNumber="320" phylum="Bryozoa" rank="species" species="zostericola">
<emphasis id="D5D1493BFF812569FC9262CA030DFA07" box="[826,1052,1484,1505]" italics="true" pageId="3" pageNumber="320">Tendra zostericola</emphasis>
</taxonomicName>
and
<emphasis id="D5D1493BFF812569FBD462CB02B0FA07" box="[1148,1441,1484,1506]" italics="true" pageId="3" pageNumber="320">Heteroecium amplectens</emphasis>
(
<bibRefCitation id="8334E8D8FF812569FCEA62ED04C8F9E7" author="Hincks T" box="[834,985,1515,1537]" pageId="3" pageNumber="320" pagination="122 - 136" refId="ref26738" refString="Hincks T. 1881. Contributions towards a general history of the marine Polyzoa. The Annals and Magazine of Natural History, 5 Series 8 (43): 1 - 14, 122 - 136." type="journal article" year="1881">Hincks, 1881</bibRefCitation>
). The first superficial description of the cellule treillissées and developing embryos inside these brood chambers was given by
<bibRefCitation id="8334E8D8FF812569FB70612E047AF9BB" author="Nordmann MA" pageId="3" pageNumber="320" pagination="185 - 191" refId="ref27386" refString="Nordmann MA. 1839. Recherches microscopiques sur l'anatomie et le developpement du Tendra zostericola, espece de polype de la section des Bryozoaires. Annales des Sciences Naturelles, Serie 2 (11): 185 - 191." type="journal article" year="1839">Nordmann (1839: 191)</bibRefCitation>
for
<taxonomicName id="20A5EEAAFF812569FC31614104FBF9BA" authorityName="NORDMANN" authorityYear="1839" box="[921,1002,1607,1628]" class="Gymnolaemata" family="Tendridae" genus="Tendra" kingdom="Animalia" order="Cheilostomatida" pageId="3" pageNumber="320" phylum="Bryozoa" rank="genus">
<emphasis id="D5D1493BFF812569FC31614104FBF9BA" box="[921,1002,1607,1628]" italics="true" pageId="3" pageNumber="320">Tendra</emphasis>
</taxonomicName>
.
<bibRefCitation id="8334E8D8FF812569FC5F614103E4F9BB" author="Repiachoff W" box="[1015,1269,1607,1629]" pageId="3" pageNumber="320" pagination="129 - 142" refId="ref27910" refString="Repiachoff W. 1875. Zur Entwickelungsgeschichte der Tendra zostericola. Zeitschrift fur Wissenschaftliche Zoologie 25: 129 - 142." type="journal article" year="1875">Repiachoff (1875: 132)</bibRefCitation>
remarked that these zooids play a role of the ovicells known in a majority of the
<taxonomicName id="20A5EEAAFF812569FC5B6182038DF97C" box="[1011,1180,1668,1690]" class="Gymnolaemata" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cheilostomata" pageId="3" pageNumber="320" phylum="Bryozoa" rank="order">Cheilostomata</taxonomicName>
. Following
<bibRefCitation id="8334E8D8FF812569FA8B618204ECF95F" author="Repiachoff W" pageId="3" pageNumber="320" pagination="129 - 142" refId="ref27910" refString="Repiachoff W. 1875. Zur Entwickelungsgeschichte der Tendra zostericola. Zeitschrift fur Wissenschaftliche Zoologie 25: 129 - 142." type="journal article" year="1875">Repiachoff, Reingard (1875)</bibRefCitation>
thought that embryos developed inside the body cavity of specialized zooids in this species. However, he believed that they could not be compared with ovicells since they possess a polypide and an ovary.
</paragraph>
<paragraph id="E71A9529FF81256EFCFB603A07AEF9BD" blockId="3.[826,1442,1392,1905]" lastBlockId="4.[142,758,1422,1904]" lastPageId="4" lastPageNumber="321" pageId="3" pageNumber="320">
<bibRefCitation id="8334E8D8FF812569FCFB603A033BF8B4" author="Ostroumoff A" box="[851,1066,1852,1874]" pageId="3" pageNumber="320" pagination="1 - 124" refId="ref27500" refString="Ostroumoff A. 1886. Die Bryozoen der Bucht von Sebastopol. Vollstandigere Ausgabe mit einem ganz neuen Theile uber die Morphologie der Bryozoen. Proceedings of the Society of Naturalists of Kazan 16 (2): 1 - 124 [in Russian]." type="journal article" year="1886">Ostroumoff (1886)</bibRefCitation>
was the first to recognize the actual position of the developing embryos in the space between the frontal membrane and the overarching spines.
<bibRefCitation id="8334E8D8FF86256EFF4E62AA0578FA24" author="Paltschikowa-Ostroumowa MW" box="[230,617,1452,1474]" pageId="4" pageNumber="321" pagination="100 - 102" refId="ref27781" refString="Paltschikowa-Ostroumowa MW. 1926. Kurze Bemerkung uber den Ovidukt bei den Bryozoen. Zoologischer Anzeiger 65: 100 - 102." type="journal article" year="1926">Paltschikowa-Ostroumowa (1926)</bibRefCitation>
and
<bibRefCitation id="8334E8D8FF86256EFD0062AA07C7FA07" author="Braiko VB" pageId="4" pageNumber="321" pagination="1119 - 1121" refId="ref25771" refString="Braiko VB. 1967. Biology of reproduction of Membranipora zostericola Nordm. (Bryozoa). Zoologicheskiy Zhurnal 46: 1119 - 1121 [in Russian]." type="journal article" year="1967">Braiko (1967)</bibRefCitation>
further studied different aspects of reproduction in
<taxonomicName id="20A5EEAAFF86256EFF7662EC06BDFA19" authorityName="Nordmann, NHM" authorityYear="1839" box="[222,428,1514,1535]" class="Gymnolaemata" family="Tendridae" genus="Tendra" kingdom="Animalia" order="Cheilostomatida" pageId="4" pageNumber="321" phylum="Bryozoa" rank="species" species="zostericola">
<emphasis id="D5D1493BFF86256EFF7662EC06BDFA19" box="[222,428,1514,1535]" italics="true" pageId="4" pageNumber="321">Tendra zostericola</emphasis>
</taxonomicName>
(see also
<bibRefCitation id="8334E8D8FF86256EFDB062EC0646F9FB" author="Occhipinti Ambrogi A &amp; d'Hondt J-L" pageId="4" pageNumber="321" pagination="191 - 198" refId="ref27426" refString="Occhipinti Ambrogi A, d'Hondt J-L. 1981. Distribution of bryozoans in brackish waters of Italy. In: Larwood GP, Nielsen C, eds. Recent and fossil Bryozoa. Fredensborg: Olsen &amp; Olsen, 191 - 198." type="book chapter" year="1981">Occhipinti Ambrogi &amp; dHondt, 1981</bibRefCitation>
, and references therein), which is known from the Mediterranean as well as the Black Sea.
</paragraph>
<caption id="B3DAC5A1FF86256EFF26639904A8FACE" ID-DOI="http://doi.org/10.5281/zenodo.5434722" ID-Zenodo-Dep="5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="4" pageNumber="321" startId="4.[142,220,1183,1202]" targetBox="[147,1416,198,1150]" targetPageId="4">
<paragraph id="E71A9529FF86256EFF26639904A8FACE" blockId="4.[142,1421,1183,1320]" pageId="4" pageNumber="321">
<emphasis id="D5D1493BFF86256EFF26639907E7FB54" bold="true" box="[142,246,1183,1202]" pageId="4" pageNumber="321">Figure 2.</emphasis>
<emphasis id="D5D1493BFF86256EFEAA63A60511FB54" box="[258,512,1183,1202]" italics="true" pageId="4" pageNumber="321">Heteroecium amplectens</emphasis>
Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm.
</paragraph>
</caption>
<paragraph id="E71A9529FF86256EFF0F61620616F896" blockId="4.[142,758,1422,1904]" pageId="4" pageNumber="321">Where several brooding zooids are present successively within the same longitudinal row, all of these zooids (possibly excepting the most distal one) will have produced their own eggs as well as brooding the embryos of the proximal neighbouring zooid. As in cheilostomes with ovicells, formation of the brood chamber by any particular zooid is presumably triggered by the development of an ovary in its proximal neighbour.</paragraph>
<paragraph id="E71A9529FF86256EFC2D628A0248FA22" blockId="4.[859,1369,1420,1508]" pageId="4" pageNumber="321">
GENUS
<emphasis id="D5D1493BFF86256EFC76628B0397FA45" box="[990,1158,1421,1444]" italics="true" pageId="4" pageNumber="321">HETEROECIUM</emphasis>
<bibRefCitation id="8334E8D8FF86256EFB26628A023FFA42" author="Hincks T" box="[1166,1326,1420,1444]" pageId="4" pageNumber="321" pagination="327 - 334" refId="ref26776" refString="Hincks T. 1892. Contributions towards a general history of the marine Polyzoa, 1880 - 91. Appendix. The Annals and Magazine of Natural History, 6 Series 9 (52): 327 - 334." type="journal article" year="1892">HINCKS, 1892</bibRefCitation>
<emphasis id="D5D1493BFF86256EFCF362AB038EFA25" box="[859,1183,1453,1476]" italics="true" pageId="4" pageNumber="321">HETEROECIUM AMPLECTENS</emphasis>
(
<bibRefCitation id="8334E8D8FF86256EFB0762AA0240FA22" author="Hincks T" box="[1199,1361,1452,1476]" pageId="4" pageNumber="321" pagination="122 - 136" refId="ref26738" refString="Hincks T. 1881. Contributions towards a general history of the marine Polyzoa. The Annals and Magazine of Natural History, 5 Series 8 (43): 1 - 14, 122 - 136." type="journal article" year="1881">HINCKS, 1881</bibRefCitation>
)
</paragraph>
<paragraph id="E71A9529FF86256EFC5B62CA03D0FA02" blockId="4.[859,1369,1420,1508]" box="[1011,1217,1484,1508]" pageId="4" pageNumber="321">
(
<figureCitation id="7F9E89ACFF86256EFC5362CA036CFA02" box="[1019,1149,1484,1508]" captionStart="Figure 2" captionStartId="4.[142,220,1183,1202]" captionTargetBox="[147,1416,198,1150]" captionTargetId="figure-344@4.[147,1418,197,1150]" captionTargetPageId="4" captionText="Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="4" pageNumber="321">FIGS 2A- D</figureCitation>
,
<figureCitation id="7F9E89ACFF86256EFB2262CA03A8FA02" box="[1162,1209,1484,1508]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="4" pageNumber="321">21B</figureCitation>
)
</paragraph>
<paragraph id="E71A9529FF86256EFC8E62FB032FF9D4" blockId="4.[806,1422,1533,1586]" pageId="4" pageNumber="321">
<emphasis id="D5D1493BFF86256EFC8E62FB049EF9F4" box="[806,911,1533,1554]" italics="true" pageId="4" pageNumber="321">Material:</emphasis>
NHM 99.5.1.702, Recent,
<collectingRegion id="25615BCBFF86256EFB6562FB0454F9D4" country="Australia" name="Western Australia" pageId="4" pageNumber="321">Western Australia</collectingRegion>
, T. Hincks Collection.
</paragraph>
<paragraph id="E71A9529FF86256EFC8E614E0314F8D2" blockId="4.[806,1422,1607,1905]" pageId="4" pageNumber="321">
<emphasis id="D5D1493BFF86256EFC8E614E04BFF9BB" box="[806,942,1608,1629]" italics="true" pageId="4" pageNumber="321">Description:</emphasis>
Non-brooding autozooids possess seven mural spines surrounding the frontal membrane. Six of these spines are short, with blunt roundish tips. The seventh spine is several times longer than the others, though approximately equal in width, and is placed on the proximal gymnocyst just behind the frontal membrane. In contrast with the other mural spines, it is basally articulated.
</paragraph>
<paragraph id="E71A9529FF86256FFC97603A067DFCC7" blockId="4.[806,1422,1607,1905]" lastBlockId="5.[162,779,196,801]" lastPageId="5" lastPageNumber="322" pageId="4" pageNumber="321">
So-called brooding zooids are actually complexes of two zooids, a proximal (maternal) autozooid (probably an autozooidal polymorph) and a distal kenozooid (
<figureCitation id="7F9E89ACFF87256FFF0267E5061AFF1F" box="[170,267,227,249]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="5" pageNumber="322">Fig. 21B</figureCitation>
). Laterally juxtaposed and flattened spines overarch the frontal wall of the kenozooid to form the brooding cavity (
<figureCitation id="7F9E89ACFF87256FFECB662606CDFED0" box="[355,476,288,310]" captionStart="Figure 2" captionStartId="4.[142,220,1183,1202]" captionTargetBox="[147,1416,198,1150]" captionTargetId="figure-344@4.[147,1418,197,1150]" captionTargetPageId="4" captionText="Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="5" pageNumber="322">Figs 2A, B</figureCitation>
,
<figureCitation id="7F9E89ACFF87256FFE4166260508FED0" box="[489,537,288,310]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="5" pageNumber="322">21B</figureCitation>
). The nonarticulated spines, numbering 1517, start their growth on the mural rim of the kenozooid and meet along the midline above the calcified frontal wall, forming a sort of medial keel. Cavities of the spines are confluent with the visceral coelom of the kenozooid (
<figureCitation id="7F9E89ACFF87256FFDD966BC05D7FE36" box="[625,710,442,464]" captionStart="Figure 2" captionStartId="4.[142,220,1183,1202]" captionTargetBox="[147,1416,198,1150]" captionTargetId="figure-344@4.[147,1418,197,1150]" captionTargetPageId="4" captionText="Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="5" pageNumber="322">Fig. 2C</figureCitation>
). The brood chamber has the shape of an elongated hemisphere, opening proximally where it is plugged by the operculum of the maternal zooid. Brood chambers are always formed in the axils of branch bifurcations, and there is never a zooid distal of the brood chamber. Communication between the maternal autozooidal polymorph and the distal kenozooid is via simple pores (
<figureCitation id="7F9E89ACFF87256FFF0265A90615FD22" box="[170,260,687,709]" captionStart="Figure 2" captionStartId="4.[142,220,1183,1202]" captionTargetBox="[147,1416,198,1150]" captionTargetId="figure-344@4.[147,1418,197,1150]" captionTargetPageId="4" captionText="Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="5" pageNumber="322">Figs 2D</figureCitation>
,
<figureCitation id="7F9E89ACFF87256FFEB965A90651FD23" box="[273,320,687,709]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="5" pageNumber="322">21B</figureCitation>
). The floor of the brood chamber is calcified, except for a peculiar area in its proximal part which is membranous with two lateral appendages (
<figureCitation id="7F9E89ACFF87256FFF02640D0632FCC6" box="[170,291,779,801]" captionStart="Figure 2" captionStartId="4.[142,220,1183,1202]" captionTargetBox="[147,1416,198,1150]" captionTargetId="figure-344@4.[147,1418,197,1150]" captionTargetPageId="4" captionText="Figure 2. Heteroecium amplectens Hincks, NHM 99.5.1.702, Recent, Western Australia. A, part of a colony with several nonbrooding zooids and one brooding zooidal complex. B, brooding zooidal complex. C, brooding zooidal complex from below (distal to the right); openings of the costae surround the floor of brood chamber. D, membranous area with two appendages in the brood-chamber floor; a communication pore in the transverse wall between the maternal zooid and distal kenozooiod can be seen in the left lower corner. Scale bars: A, C = 100 Mm; B, D = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5434722" httpUri="https://zenodo.org/record/5434722/files/figure.png" pageId="5" pageNumber="322">Figs 2C, D</figureCitation>
,
<figureCitation id="7F9E89ACFF87256FFE87640D064EFCC7" box="[303,351,779,801]" captionStart="Figure 21" captionStartId="35.[162,240,1239,1258]" captionTargetBox="[328,1279,202,1207]" captionTargetId="figure-89@35.[320,1281,197,1203]" captionTargetPageId="35" captionText="Figure 21. Schematic diagrams of brood chambers in Tendridae (A, B) and Calloporidae (C-E) in longitudinal and transverse section, showing maternal and distal zooids (fossil spinose ovicells reconstructed). A, Tendra zostericola. B, Heteroecium sp. C, Distelopora bipilata and D. langi. D, Distelopora spinifera, Unidistelopora krauseae. E, Gilbertopora larwoodi." figureDoi="http://doi.org/10.5281/zenodo.5434760" httpUri="https://zenodo.org/record/5434760/files/figure.png" pageId="5" pageNumber="322">21B</figureCitation>
).
</paragraph>
<paragraph id="E71A9529FF87256FFF0A644E0515F8F2" blockId="5.[162,779,840,1905]" pageId="5" pageNumber="322">
<emphasis id="D5D1493BFF87256FFF0A644E061CFCBB" box="[162,269,840,861]" italics="true" pageId="5" pageNumber="322">Remarks:</emphasis>
The first description of this species was given by
<bibRefCitation id="8334E8D8FF87256FFEA7646106A0FC9B" author="Hincks T" box="[271,433,871,893]" pageId="5" pageNumber="322" pagination="122 - 136" refId="ref26738" refString="Hincks T. 1881. Contributions towards a general history of the marine Polyzoa. The Annals and Magazine of Natural History, 5 Series 8 (43): 1 - 14, 122 - 136." type="journal article" year="1881">Hincks (1881)</bibRefCitation>
who correctly recognized the spinose structures as brood chambers. However, he believed them to be single zooids with the upper portion of the zooecial aperture... much extended and roofed in by a number of (soldered) marginal spines. Hincks also wrote that the zooecium... is divided into two chambers one for the polypide, the other for the embryo (1881: 130). In a later work,
<bibRefCitation id="8334E8D8FF87256FFDC46338041BFBB2" author="Hincks T" box="[620,778,1086,1108]" pageId="5" pageNumber="322" pagination="327 - 334" refId="ref26776" refString="Hincks T. 1892. Contributions towards a general history of the marine Polyzoa, 1880 - 91. Appendix. The Annals and Magazine of Natural History, 6 Series 9 (52): 327 - 334." type="journal article" year="1892">Hincks (1892)</bibRefCitation>
introduced the genus
<emphasis id="D5D1493BFF87256FFE0F635B0524FB94" box="[423,565,1117,1138]" italics="true" pageId="5" pageNumber="322">Heteroecium</emphasis>
and repeated the description mentioned above.
<bibRefCitation id="8334E8D8FF87256FFDA6637D041BFB77" author="Levinsen GMR" box="[526,778,1147,1169]" pageId="5" pageNumber="322" refId="ref27042" refString="Levinsen GMR. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa. Copenhagen: Bagge F." type="book" year="1909">Levinsen (1909: 148)</bibRefCitation>
described a second species,
<taxonomicName id="20A5EEAAFF87256FFE57639C0586FB49" box="[511,663,1178,1199]" pageId="5" pageNumber="322" rank="species" species="brevispina">
<emphasis id="D5D1493BFF87256FFE57639C0586FB49" box="[511,663,1178,1199]" italics="true" pageId="5" pageNumber="322">H. brevispina</emphasis>
</taxonomicName>
(as Var.
<taxonomicName id="20A5EEAAFF87256FFF0A63BE0608FB2B" box="[162,281,1208,1229]" pageId="5" pageNumber="322" rank="species" species="brevispina">
<emphasis id="D5D1493BFF87256FFF0A63BE0608FB2B" box="[162,281,1208,1229]" italics="true" pageId="5" pageNumber="322">brevispina</emphasis>
</taxonomicName>
n.), with some clear differences (larger size, more mural spines, shorter proximal spine, etc.) but a brood chamber having the same structure. Not surprisingly,
<bibRefCitation id="8334E8D8FF87256FFE9F62120535FACC" author="Levinsen GMR" box="[311,548,1300,1322]" pageId="5" pageNumber="322" refId="ref27042" refString="Levinsen GMR. 1909. Morphological and systematic studies on the cheilostomatous Bryozoa. Copenhagen: Bagge F." type="book" year="1909">Levinsen (1909: 148)</bibRefCitation>
did not understand the significance of the situation of the aperture behind the area formed by the spines, since he thought that the brood chamber consisted of a single zooid. Nevertheless, he correctly understood the structure of the brooding cavity, and, following
<bibRefCitation id="8334E8D8FF87256FFD8162A80417FA22" author="Ostroumoff A" box="[553,774,1454,1476]" pageId="5" pageNumber="322" pagination="1 - 124" refId="ref27500" refString="Ostroumoff A. 1886. Die Bryozoen der Bucht von Sebastopol. Vollstandigere Ausgabe mit einem ganz neuen Theile uber die Morphologie der Bryozoen. Proceedings of the Society of Naturalists of Kazan 16 (2): 1 - 124 [in Russian]." type="journal article" year="1886">Ostroumoff (1886)</bibRefCitation>
, wrote that the acanthostegal ooecia... are cavities internally limited by the covering membrane of the zooecium and externally by a cover made up of two rows of hollow spines (
<bibRefCitation id="8334E8D8FF87256FFE15612E058FF9D8" author="Levinsen GMR" box="[445,670,1576,1598]" pageId="5" pageNumber="322" pagination="1 - 31" refId="ref27019" refString="Levinsen GMR. 1902. Studies on Bryozoa. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn 54: 1 - 31." type="journal article" year="1902">Levinsen, 1902: 17</bibRefCitation>
). Images of
<taxonomicName id="20A5EEAAFF87256FFF6E614E064FF9BA" box="[198,350,1607,1629]" pageId="5" pageNumber="322" rank="species" species="brevispina">
<emphasis id="D5D1493BFF87256FFF6E614E064FF9BA" box="[198,350,1607,1629]" italics="true" pageId="5" pageNumber="322">H. brevispina</emphasis>
</taxonomicName>
recently placed by Dr P. E. Bock on the Bryozoa Home Page website (http://www. civgeo.rmit.edu.au/bryozoa/cheilostomata/tendridae/ heteamp.html) show the brood chamber to be constructed of 1213 flattened, nonarticulated spines. Closure of the brood-chamber opening by the zooidal operculum is also clearly seen.
</paragraph>
<paragraph id="E71A9529FF87256FFF1360180336FEFE" blockId="5.[162,779,840,1905]" lastBlockId="5.[826,1441,196,280]" pageId="5" pageNumber="322">The unusual membranous area in the floor of the brood chamber is interpreted as a rudiment of the frontal membrane to which are attached the parietal muscles involved in tentacle crown eversion in the autozooids. Sections of decalcified material would help to resolve this issue.</paragraph>
</subSubSection>
</treatment>
</document>