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<mods:title id="949726E8C3CE816C5C19A0A74FABA499">Four new species and a ribosomal phylogeny of Rhabdopleura (Hemichordata: Graptolithina) from New Zealand, with a review and key to all described extant taxa</mods:title>
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<paragraph id="8B97A15BFFCEB95EEAF0FAC2F634FACB" blockId="11.[151,854,1326,1353]" box="[151,854,1326,1353]" pageId="11" pageNumber="334">
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<taxonomicName id="4C28DAD8FFCEB95EEAF0FAC2F702FACB" ID-CoL="4RW6V" authority="Jullien, 1890" authorityName="Jullien" authorityYear="1890" box="[151,608,1326,1353]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="11" pageNumber="334" phylum="Hemichordata" rank="species" species="grimaldii">
<emphasis id="B95C7D49FFCEB95EEAF0FAC2F4D8FACA" bold="true" box="[151,442,1326,1353]" italics="true" pageId="11" pageNumber="334">Rhabdopleura grimaldii</emphasis>
<bibRefCitation id="EFB9DCAAFFCEB95EEBA6FAC2F702FACB" author="Jullien, J." box="[449,608,1326,1352]" pageId="11" pageNumber="334" pagination="180 - 183" refId="ref22649" refString="Jullien, J. (1890) Description d'un Bryozoaire nouveau du genre Rhabdopleura. Bulletin de la Societe Zoologique de France, 15, 180 - 183. https: // doi. org / 10.5962 / bhl. part. 18723" type="journal article" year="1890">Jullien, 1890</bibRefCitation>
</taxonomicName>
(
<figureCitation id="1313BDDEFFCEB95EE808FAC2F795FACB" box="[623,759,1326,1352]" captionStart="FIGURE 4" captionStartId="10.[151,250,1795,1819]" captionTargetBox="[175,1413,182,1758]" captionTargetId="figure-28@10.[162,1426,181,1771]" captionTargetPageId="10" captionText="FIGURE 4. Original illustrations of described species of Rhabdopleura characterized by indirect inception of ringed erect tubes from creeping tubes via an intervening adnate portion that comprises a blind-ending branch. A, Rhabdopleura normani (from Allman 1869, pl. 8, fig. 1). Note that most of the colony is adherent; only the ringed tubes (rt) are erect, the proximal part of each side branch (ppsb) is adherent and the oblique sutures are not shown. B, schematic interpretative depiction of R. normani (after Andres 1977). C, D, Rhabdopleura grimaldii (from Jullien 1890, p. 181); note that ringed erect tubes (rt) are mostly lacking, with only the adherent proximal parts of side branches (ppsb) present. D is a cropped part of C to show the proximal parts of two side branches, with fusellar sutures and Julliens putative very fine tubes (vft)—actually the side walls of the tapering pectocaulus, the proximal end of which is transparent, hence the apparent gap between it and the pectocaulus of the creeping tube. E, F, Rhabdopleura striata (from Schepotieff 1909, pl. 7, figs 2, 7); E shows longitudinally striated erect parts of side branches; oblique sutures on prostrate parts omitted in E but shown in F, with characteristic web of fusellar sutures (wfs) in branch angles from main creeping tube. G, Rhabdopleura manubialis (from Jullien &amp; Calvet, 1903, pl. 1, fig. 2) with dormant bodies (db)." figureDoi="http://doi.org/10.5281/zenodo.10821369" httpUri="https://zenodo.org/record/10821369/files/figure.png" pageId="11" pageNumber="334">Figs 4C, D</figureCitation>
;
<figureCitation id="1313BDDEFFCEB95EE962FAC2F62CFACB" box="[773,846,1326,1352]" captionStart="FIGURE 6" captionStartId="13.[151,250,1294,1318]" captionTargetBox="[262,1325,181,1270]" captionTargetId="figure-17@13.[249,1337,181,1270]" captionTargetPageId="13" captionText="FIGURE 6. Rhabdopleura grimaldii Jullien, 1890 (AD, syntype MNHN-IB-2014-386) and R. manubialis Jullien &amp; Calvet, 1903 (E, syntype MNHN-IB-2014-387). A, B, oblique views of two erect ringed tubes. C, adherent proximal part of a side branch (apsb), with a tapering pectocaulus (tp) (its side walls appearing as converging brown lines) and the broken base of an erect tube (bbet). D, similar to C, but also showing the creeping tube (ct) from which the broader side branch originated; note the faint outlines of oblique sutures on the creeping tube and side branch. E, erect ringed tube. Scale bars: A, B, 100 μm; CE, 200 μm. Images cropped from photos supplied by Pierre Lozuet, MNHN, Paris." figureDoi="http://doi.org/10.5281/zenodo.10821375" httpUri="https://zenodo.org/record/10821375/files/figure.png" pageId="11" pageNumber="334">6AD</figureCitation>
)
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<materialsCitation id="3B40AB06FFCEB95EEAF0FA9AF1CAFA37" county="Azores" elevation="318" latitude="38.4" location="Off" longLatPrecision="6" longitude="-28.0239" municipality="Pico Island" pageId="11" pageNumber="334" specimenCount="1" typeStatus="holotype">
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locality.
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<location id="8EF7F780FFCEB95EEB27FA9AF404FA13" LSID="urn:lsid:plazi:treatment:0381104DFFCEB95BEAF0FAC2F0E9FE52:8EF7F780FFCEB95EEB27FA9AF404FA13" box="[320,358,1398,1424]" county="Azores" latitude="38.4" longLatPrecision="6" longitude="-28.0239" municipality="Pico Island" name="Off" pageId="11" pageNumber="334">Off</location>
the southeastern tip of
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,
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, on dead shell and bryozoans [
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,
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in
<bibRefCitation id="EFB9DCAAFFCEB95EEB25FA76F49FFA37" author="Stebbing, A. R. D." box="[322,509,1434,1460]" pageId="11" pageNumber="334" pagination="209 - 221" refId="ref25339" refString="Stebbing, A. R. D. (1970 a) The status and ecology of Rhabdopleura compacta (Hemichordata) from Plymouth. Journal of the Marine Biological Association of the United Kingdom, 50, 209 - 221. https: // doi. org / 10.1017 / S 0025315400000722" type="journal article" year="1970">Stebbing (1970a</bibRefCitation>
, p. 211), the original Paris meridian corrected to Greenwich]
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.
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<subSubSection id="C332F2D0FFCEB95EEAA0FA52F7FFF8E7" pageId="11" pageNumber="334" type="key">
<paragraph id="8B97A15BFFCEB95EEAA0FA52F7FFF8E7" blockId="11.[151,1437,1398,2037]" pageId="11" pageNumber="334">
<emphasis id="B95C7D49FFCEB95EEAA0FA52F405FA5B" bold="true" box="[199,359,1470,1496]" pageId="11" pageNumber="334">Key features.</emphasis>
Inception of ringed erect tubes is indirect.
<bibRefCitation id="EFB9DCAAFFCEB95EE934FA52F693FA5B" author="Jullien, J." box="[851,1009,1470,1496]" pageId="11" pageNumber="334" pagination="180 - 183" refId="ref22649" refString="Jullien, J. (1890) Description d'un Bryozoaire nouveau du genre Rhabdopleura. Bulletin de la Societe Zoologique de France, 15, 180 - 183. https: // doi. org / 10.5962 / bhl. part. 18723" type="journal article" year="1890">Jullien (1890)</bibRefCitation>
described the 12 cm-long colony as consisting of a primary bifurcating creeping tube (stolon) containing the pectocaulus (filet chitineux), with short blind side branches on either side, each comprising an adherent portion (zooecium) with zigzag sutures, terminating in an erect péristomie, with annular fusellar collars. This arrangement corresponds to that in
<taxonomicName id="4C28DAD8FFCEB95EEEC4F9C7F042F9C7" ID-CoL="4RW6Y" box="[1187,1312,1579,1604]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="11" pageNumber="334" phylum="Hemichordata" rank="species" species="normani">
<emphasis id="B95C7D49FFCEB95EEEC4F9C7F042F9C7" box="[1187,1312,1579,1604]" italics="true" pageId="11" pageNumber="334">R. normani</emphasis>
</taxonomicName>
,
<emphasis id="B95C7D49FFCEB95EEF4AF9C7F02DF9C7" box="[1325,1359,1579,1604]" italics="true" pageId="11" pageNumber="334">i.e.</emphasis>
zooidcontaining side tubes are proximally adherent before bending at an angle upwards. Zooecia are said to be a little wider than the stolon. The main distinguishing character of this species was described as two very thin tubes (tubes très fins) in the adherent proximal parts of the blind side branches, reflected externally as two dark-brown lines (lignes brun foncé), the distal ends of which converge without meeting at the proximal end of the péristomie (
<figureCitation id="1313BDDEFFCEB95EEF08F956F582F97B" captionStart="FIGURE 6" captionStartId="13.[151,250,1294,1318]" captionTargetBox="[262,1325,181,1270]" captionTargetId="figure-17@13.[249,1337,181,1270]" captionTargetPageId="13" captionText="FIGURE 6. Rhabdopleura grimaldii Jullien, 1890 (AD, syntype MNHN-IB-2014-386) and R. manubialis Jullien &amp; Calvet, 1903 (E, syntype MNHN-IB-2014-387). A, B, oblique views of two erect ringed tubes. C, adherent proximal part of a side branch (apsb), with a tapering pectocaulus (tp) (its side walls appearing as converging brown lines) and the broken base of an erect tube (bbet). D, similar to C, but also showing the creeping tube (ct) from which the broader side branch originated; note the faint outlines of oblique sutures on the creeping tube and side branch. E, erect ringed tube. Scale bars: A, B, 100 μm; CE, 200 μm. Images cropped from photos supplied by Pierre Lozuet, MNHN, Paris." figureDoi="http://doi.org/10.5281/zenodo.10821375" httpUri="https://zenodo.org/record/10821375/files/figure.png" pageId="11" pageNumber="334">Fig. 6C, D</figureCitation>
). Oblique sutures with zigzag angles were described as restricted to the zooecium; the primary creeping tube itself was dépourvue de stries (free of streaks), with zooids retracting into the adherent portion of the side branches as in
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<emphasis id="B95C7D49FFCEB95EEB5BF8CBF4D8F8C3" box="[316,442,1831,1856]" italics="true" pageId="11" pageNumber="334">R. normani</emphasis>
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. Annulated erect tubes were mostly missing from his material; some are broken and only one appears intact in his drawing (
<figureCitation id="1313BDDEFFCEB95EE87FF8A6F70DF8E7" box="[536,623,1866,1892]" captionStart="FIGURE 4" captionStartId="10.[151,250,1795,1819]" captionTargetBox="[175,1413,182,1758]" captionTargetId="figure-28@10.[162,1426,181,1771]" captionTargetPageId="10" captionText="FIGURE 4. Original illustrations of described species of Rhabdopleura characterized by indirect inception of ringed erect tubes from creeping tubes via an intervening adnate portion that comprises a blind-ending branch. A, Rhabdopleura normani (from Allman 1869, pl. 8, fig. 1). Note that most of the colony is adherent; only the ringed tubes (rt) are erect, the proximal part of each side branch (ppsb) is adherent and the oblique sutures are not shown. B, schematic interpretative depiction of R. normani (after Andres 1977). C, D, Rhabdopleura grimaldii (from Jullien 1890, p. 181); note that ringed erect tubes (rt) are mostly lacking, with only the adherent proximal parts of side branches (ppsb) present. D is a cropped part of C to show the proximal parts of two side branches, with fusellar sutures and Julliens putative very fine tubes (vft)—actually the side walls of the tapering pectocaulus, the proximal end of which is transparent, hence the apparent gap between it and the pectocaulus of the creeping tube. E, F, Rhabdopleura striata (from Schepotieff 1909, pl. 7, figs 2, 7); E shows longitudinally striated erect parts of side branches; oblique sutures on prostrate parts omitted in E but shown in F, with characteristic web of fusellar sutures (wfs) in branch angles from main creeping tube. G, Rhabdopleura manubialis (from Jullien &amp; Calvet, 1903, pl. 1, fig. 2) with dormant bodies (db)." figureDoi="http://doi.org/10.5281/zenodo.10821369" httpUri="https://zenodo.org/record/10821369/files/figure.png" pageId="11" pageNumber="334">Fig. 4C</figureCitation>
, rt).
</paragraph>
</subSubSection>
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<emphasis id="B95C7D49FFCEB95EEAA0F882F421F80B" bold="true" box="[199,323,1902,1928]" pageId="11" pageNumber="334">Comment.</emphasis>
<materialsCitation id="3B40AB06FFCEB959EB2CF882F4DCFF7A" county="The Museum National d'Histoire Naturelle" lastPageId="12" lastPageNumber="335" location="Dr Pierre Lozuet" municipality="Paris" pageId="11" pageNumber="334" specimenCount="1" stateProvince="Monaghan" typeStatus="syntype">
<collectingCounty id="62F6D9D7FFCEB95EEB2CF882F64DF80B" box="[331,815,1902,1928]" pageId="11" pageNumber="334">The Muséum National dHistoire Naturelle</collectingCounty>
(
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),
<collectingMunicipality id="6BF33B21FFCEB95EE9C9F882F68AF80B" box="[942,1000,1902,1928]" pageId="11" pageNumber="334">Paris</collectingMunicipality>
, was contacted in the hope that photos could be made of the species (MNHN-IB-2014-386).
<location id="8EF7F780FFCEB95EE88EF87FF6C1F82E" LSID="urn:lsid:plazi:treatment:0381104DFFCEB95BEAF0FAC2F0E9FE52:8EF7F780FFCEB95EE88EF87FF6C1F82E" box="[745,931,1938,1965]" county="The Museum National d'Histoire Naturelle" municipality="Paris" name="Dr Pierre Lozuet" pageId="11" pageNumber="334" stateProvince="Monaghan">Dr Pierre Lozuet</location>
of the
<location id="8EF7F780FFCEB95EE997F87EF19AF82E" LSID="urn:lsid:plazi:treatment:0381104DFFCEB95BEAF0FAC2F0E9FE52:8EF7F780FFCEB95EE997F87EF19AF82E" box="[1008,1272,1938,1965]" county="The Museum National d'Histoire Naturelle" municipality="Paris" name="Collections Department" pageId="11" pageNumber="334" stateProvince="Monaghan">Collections Department</location>
(
<location id="8EF7F780FFCEB95EEF61F87EF475F853" LSID="urn:lsid:plazi:treatment:0381104DFFCEB95BEAF0FAC2F0E9FE52:8EF7F780FFCEB95EEF61F87EF475F853" county="The Museum National d'Histoire Naturelle" municipality="Paris" name="Direction des Collections" pageId="11" pageNumber="334" stateProvince="Monaghan">Direction des Collections</location>
) sent reflected-light images, which included two high-resolution close-ups, with scale bars.
<location id="8EF7F780FFCEB95EEF79F85AF02BF853" LSID="urn:lsid:plazi:treatment:0381104DFFCEB95BEAF0FAC2F0E9FE52:8EF7F780FFCEB95EEF79F85AF02BF853" box="[1310,1353,1974,2000]" county="The Museum National d'Histoire Naturelle" municipality="Paris" name="The" pageId="11" pageNumber="334" stateProvince="Monaghan">The</location>
unique
<typeStatus id="54931FF9FFCEB95EEAF0F837F58FF876" box="[151,237,2011,2037]" pageId="11" pageNumber="334" type="syntype">syntype</typeStatus>
is on a worm tube. It is dry and the overall colony seems to bear little resemblance to Julliens original illustration. Additionally, most of the creeping tubes are highly transparent and reflective, rendering details like fusellar sutures problematic. Notwithstanding, the images yielded enough information to discern key features and to make some measurements
</materialsCitation>
.
</paragraph>
<caption id="DF57F1D3FFC9B959EAF0F8C3F128F85B" ID-DOI="http://doi.org/10.5281/zenodo.10821371" ID-Zenodo-Dep="10821371" httpUri="https://zenodo.org/record/10821371/files/figure.png" pageId="12" pageNumber="335" startId="12.[151,250,1839,1863]" targetBox="[201,1388,286,1803]" targetPageId="12" targetType="figure">
<paragraph id="8B97A15BFFC9B959EAF0F8C3F128F85B" blockId="12.[151,1436,1839,2008]" pageId="12" pageNumber="335">
<emphasis id="B95C7D49FFC9B959EAF0F8C3F470F8CB" bold="true" box="[151,274,1839,1864]" pageId="12" pageNumber="335">FIGURE 5.</emphasis>
Original illustrations of three described species of
<taxonomicName id="4C28DAD8FFC9B959E976F8C3F6C0F8C4" authorityName="Allman" authorityYear="1869" box="[785,930,1839,1863]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="genus">
<emphasis id="B95C7D49FFC9B959E976F8C3F6C0F8C4" box="[785,930,1839,1863]" italics="true" pageId="12" pageNumber="335">Rhabdopleura</emphasis>
</taxonomicName>
characterized by direct-frontal inception of ringed erect tubes (AC). Erect-tube inception in
<taxonomicName id="4C28DAD8FFC9B959E82AF8B8F7AEF8E8" authorityName="Hincks" authorityYear="1880" box="[589,716,1876,1899]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="species" species="compacta">
<emphasis id="B95C7D49FFC9B959E82AF8B8F7AEF8E8" box="[589,716,1876,1899]" italics="true" pageId="12" pageNumber="335">R. compacta</emphasis>
</taxonomicName>
(DF) appears to be a form of indirect erect-tube inception. Note that the direction of zooid budding is the same in A to C.
<emphasis id="B95C7D49FFC9B959E8C0F89BF7DDF80C" bold="true" box="[679,703,1911,1935]" pageId="12" pageNumber="335">A,</emphasis>
<taxonomicName id="4C28DAD8FFC9B959E8A2F89BF6DAF80C" baseAuthorityName="Sars" baseAuthorityYear="1872" box="[709,952,1911,1935]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="species" species="mirabilis">
<emphasis id="B95C7D49FFC9B959E8A2F89BF6DAF80C" box="[709,952,1911,1935]" italics="true" pageId="12" pageNumber="335">Rhabdopleura mirabilis</emphasis>
</taxonomicName>
(from
<bibRefCitation id="EFB9DCAAFFC9B959E99AF894F101F813" author="Sars, G. O." box="[1021,1123,1912,1936]" pageId="12" pageNumber="335" refId="ref24443" refString="Sars, G. O. (1872) On some remarkable forms of animal life from the great deeps off the Norwegian coast I. Partly from posthumous manuscripts of the late Professor Dr. Michael Sars. University-Program for the 1 st half-year 1869. Brogger &amp; Christie, Cristiania, 82 pp., 6 pls. https: // doi. org / 10.5962 / bhl. title. 8861" type="book" year="1872">Sars 1872</bibRefCitation>
, pl. 1, fig. 5).
<emphasis id="B95C7D49FFC9B959EE89F894F067F813" bold="true" box="[1262,1285,1912,1936]" pageId="12" pageNumber="335">B,</emphasis>
<taxonomicName id="4C28DAD8FFC9B959EF6CF89BF593F830" authorityName="Norman" authorityYear="1921" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="species" species="annulata">
<emphasis id="B95C7D49FFC9B959EF6CF89BF593F830" italics="true" pageId="12" pageNumber="335">Rhabdopleura annulata</emphasis>
</taxonomicName>
(from
<bibRefCitation id="EFB9DCAAFFC9B959EB51F870F4A1F837" author="Norman, J. R." box="[310,451,1948,1972]" pageId="12" pageNumber="335" pagination="95 - 102" refId="ref23857" refString="Norman, J. R. (1921) Rhabdopleura. British Antarctic &quot; Terra Nova &quot; Expedition Natural History Reports, Zoology, 4, 95 - 102." type="journal article" year="1921">Norman 1921</bibRefCitation>
, fig. 4).
<emphasis id="B95C7D49FFC9B959E871F877F74DF830" bold="true" box="[534,559,1947,1971]" pageId="12" pageNumber="335">C,</emphasis>
<taxonomicName id="4C28DAD8FFC9B959E853F877F648F830" authorityName="Beli, Cameron &amp; Piraino" authorityYear="2018" box="[564,810,1947,1971]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="species" species="recondita">
<emphasis id="B95C7D49FFC9B959E853F877F648F830" box="[564,810,1947,1971]" italics="true" pageId="12" pageNumber="335">Rhabdopleura recondita</emphasis>
</taxonomicName>
(after
<bibRefCitation id="EFB9DCAAFFC9B959E90BF877F16AF837" author="Beli, E. &amp; Aglieri, G. &amp; Strano, F. &amp; Maggioni, D. &amp; Telford, M. &amp; Piraino, S. &amp; Cameron, C. B." box="[876,1032,1947,1972]" pageId="12" pageNumber="335" pagination="100 - 110" refId="ref20981" refString="Beli, E., Aglieri, G., Strano, F., Maggioni, D., Telford, M., Piraino, S. &amp; Cameron, C. B. (2018) The zoogeography of extant rhabdopleurid hemichordates (Pterobranchia: Graptolithina), with a new species from the Mediterranean Sea. Invertebrate Systematics, 32, 100 - 110. https: // doi. org / 10.1071 / IS 17021" type="journal article" year="2018">
Beli
<emphasis id="B95C7D49FFC9B959E9FBF870F6AAF830" box="[924,968,1947,1971]" italics="true" pageId="12" pageNumber="335">et al</emphasis>
. 2018
</bibRefCitation>
, fig. 2C).
<emphasis id="B95C7D49FFC9B959EE0CF870F1FCF837" bold="true" box="[1131,1182,1948,1972]" pageId="12" pageNumber="335">DF,</emphasis>
<taxonomicName id="4C28DAD8FFC9B959EEC4F877F0FEF830" authorityName="Hincks" authorityYear="1880" box="[1187,1436,1947,1971]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="12" pageNumber="335" phylum="Hemichordata" rank="species" species="compacta">
<emphasis id="B95C7D49FFC9B959EEC4F877F0FEF830" box="[1187,1436,1947,1971]" italics="true" pageId="12" pageNumber="335">Rhabdopleura compacta</emphasis>
</taxonomicName>
(respectively after
<bibRefCitation id="EFB9DCAAFFC9B959EB32F853F4BAF85B" author="Hincks, T." box="[341,472,1983,2008]" pageId="12" pageNumber="335" refId="ref22515" refString="Hincks, T. (1880) A History of the British Marine Polyzoa. Vols. 1 &amp; 2. Van Voorst, London, cxli + 601 pp. &amp; 83 pls." type="book" year="1880">Hincks 1880</bibRefCitation>
, pl. 72, fig. 8;
<bibRefCitation id="EFB9DCAAFFC9B959E80BF853F66CF854" author="Stebbing, A. R. D." box="[620,782,1983,2007]" pageId="12" pageNumber="335" pagination="205 - 212" refId="ref25392" refString="Stebbing, A. R. D. (1970 b) Aspects of the reproduction and life cycle of Rhabdopleura compacta (Hemichordata). Marine Biology, 5, 205 - 212. https: // doi. org / 10.1007 / bf 00346908" type="journal article" year="1970">Stebbing 1970b</bibRefCitation>
, fig. 1;
<bibRefCitation id="EFB9DCAAFFC9B959E93CF853F699F85B" author="Stebbing, A. R. D." box="[859,1019,1983,2008]" pageId="12" pageNumber="335" pagination="209 - 221" refId="ref25339" refString="Stebbing, A. R. D. (1970 a) The status and ecology of Rhabdopleura compacta (Hemichordata) from Plymouth. Journal of the Marine Biological Association of the United Kingdom, 50, 209 - 221. https: // doi. org / 10.1017 / S 0025315400000722" type="journal article" year="1970">Stebbing 1970a</bibRefCitation>
, fig. 3).
</paragraph>
</caption>
<caption id="DF57F1D3FFC8B958EAF0FAE2F6A2FA59" ID-DOI="http://doi.org/10.5281/zenodo.10821375" ID-Zenodo-Dep="10821375" httpUri="https://zenodo.org/record/10821375/files/figure.png" pageId="13" pageNumber="336" startId="13.[151,250,1294,1318]" targetBox="[262,1325,181,1270]" targetPageId="13" targetType="figure">
<paragraph id="8B97A15BFFC8B958EAF0FAE2F6A2FA59" blockId="13.[151,1437,1294,1499]" pageId="13" pageNumber="336">
<emphasis id="B95C7D49FFC8B958EAF0FAE2F476FAA5" bold="true" box="[151,276,1294,1318]" pageId="13" pageNumber="336">FIGURE 6.</emphasis>
<taxonomicName id="4C28DAD8FFC8B958EB7BFAE2F7FCFAA5" authority="Jullien, 1890" authorityName="Jullien" authorityYear="1890" box="[284,670,1294,1318]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="13" pageNumber="336" phylum="Hemichordata" rank="species" species="grimaldii">
<emphasis id="B95C7D49FFC8B958EB7BFAE2F770FAA5" box="[284,530,1294,1318]" italics="true" pageId="13" pageNumber="336">Rhabdopleura grimaldii</emphasis>
<bibRefCitation id="EFB9DCAAFFC8B958E87DFAE2F7FCFAA5" author="Jullien, J." box="[538,670,1294,1318]" pageId="13" pageNumber="336" pagination="180 - 183" refId="ref22649" refString="Jullien, J. (1890) Description d'un Bryozoaire nouveau du genre Rhabdopleura. Bulletin de la Societe Zoologique de France, 15, 180 - 183. https: // doi. org / 10.5962 / bhl. part. 18723" type="journal article" year="1890">Jullien, 1890</bibRefCitation>
</taxonomicName>
(AD, syntype MNHN-IB-2014-386) and
<taxonomicName id="4C28DAD8FFC8B958EE3FFAE3F5A8FAC9" authority="Jullien &amp; Calvet, 1903" authorityName="Jullien &amp; Calvet" authorityYear="1903" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="13" pageNumber="336" phylum="Hemichordata" rank="species" species="manubialis">
<emphasis id="B95C7D49FFC8B958EE3FFAE3F184FAA5" box="[1112,1254,1294,1318]" italics="true" pageId="13" pageNumber="336">R. manubialis</emphasis>
<bibRefCitation id="EFB9DCAAFFC8B958EE8AFAE2F5A8FAC9" author="Jullien, J. &amp; Calvet, L." pageId="13" pageNumber="336" pagination="1 - 188" refId="ref22692" refString="Jullien, J. &amp; Calvet, L. (1903) Bryozoaires provenant des campagnes de l'Hirondelle. Resultats des Campagnes scientifiques accompli par le Prince Albert I, 23, 1 - 188, 18 pls." type="journal article" year="1903">Jullien &amp; Calvet, 1903</bibRefCitation>
</taxonomicName>
(E, syntype MNHN-IB-2014-387).
<emphasis id="B95C7D49FFC8B958E826FADEF71BFAC9" bold="true" box="[577,633,1330,1354]" pageId="13" pageNumber="336">A, B,</emphasis>
oblique views of two erect ringed tubes.
<emphasis id="B95C7D49FFC8B958EE4AFADEF124FAC9" bold="true" box="[1069,1094,1330,1354]" pageId="13" pageNumber="336">C,</emphasis>
adherent proximal part of a side branch (apsb), with a tapering pectocaulus (tp) (its side walls appearing as converging brown lines) and the broken base of an erect tube (bbet).
<emphasis id="B95C7D49FFC8B958EB2BFA96F406FA11" bold="true" box="[332,356,1402,1426]" pageId="13" pageNumber="336">D,</emphasis>
similar to C, but also showing the creeping tube (ct) from which the broader side branch originated; note the faint outlines of oblique sutures on the creeping tube and side branch.
<emphasis id="B95C7D49FFC8B958E9E0FA72F6FCFA35" bold="true" box="[903,926,1438,1462]" pageId="13" pageNumber="336">E,</emphasis>
erect ringed tube. Scale bars: A, B, 100 μm; CE, 200 μm. Images cropped from photos supplied by Pierre Lozuet, MNHN, Paris.
</paragraph>
</caption>
<paragraph id="8B97A15BFFC8B95BEAA0F9C0F710FE9E" blockId="13.[151,1437,1580,2039]" lastBlockId="14.[151,1437,151,465]" lastPageId="14" lastPageNumber="337" pageId="13" pageNumber="336">
Primary creeping tubes (convexity) are 146167 (158) μm wide (
<emphasis id="B95C7D49FFC8B958E9D1F9C1F6A6F9C5" box="[950,964,1581,1606]" italics="true" pageId="13" pageNumber="336">n</emphasis>
= 5), with pectocaulus width 2144 (31) μm (
<emphasis id="B95C7D49FFC8B958EAAAF9BDF5B9F9E9" box="[205,219,1617,1642]" italics="true" pageId="13" pageNumber="336">n</emphasis>
= 8). Contra Jullien, primary creeping tubes have surface fusellar sutures; a sequence of nine point-to-point zigzag angles was measured, ranging from 6376°, with mean 70° and mode 65°. In contrast, the adherent parts of blind side branches have a higher mean convexity width [134201 (177 μm),
<emphasis id="B95C7D49FFC8B958EE6CF975F17BF931" box="[1035,1049,1689,1714]" italics="true" pageId="13" pageNumber="336">n</emphasis>
= 5]. The most striking character, which is unique to this species, is the pectocaulus of an adherent side branch. It is defined by a pair of thin brown lines (its side walls) that converge toward the base of the erect ringed tube (
<figureCitation id="1313BDDEFFC8B958E980F90CF100F978" box="[999,1122,1760,1787]" captionStart="FIGURE 6" captionStartId="13.[151,250,1294,1318]" captionTargetBox="[262,1325,181,1270]" captionTargetId="figure-17@13.[249,1337,181,1270]" captionTargetPageId="13" captionText="FIGURE 6. Rhabdopleura grimaldii Jullien, 1890 (AD, syntype MNHN-IB-2014-386) and R. manubialis Jullien &amp; Calvet, 1903 (E, syntype MNHN-IB-2014-387). A, B, oblique views of two erect ringed tubes. C, adherent proximal part of a side branch (apsb), with a tapering pectocaulus (tp) (its side walls appearing as converging brown lines) and the broken base of an erect tube (bbet). D, similar to C, but also showing the creeping tube (ct) from which the broader side branch originated; note the faint outlines of oblique sutures on the creeping tube and side branch. E, erect ringed tube. Scale bars: A, B, 100 μm; CE, 200 μm. Images cropped from photos supplied by Pierre Lozuet, MNHN, Paris." figureDoi="http://doi.org/10.5281/zenodo.10821375" httpUri="https://zenodo.org/record/10821375/files/figure.png" pageId="13" pageNumber="336">Fig. 6C, D</figureCitation>
). Because the central part is thin-walled and clear,
<bibRefCitation id="EFB9DCAAFFC8B958EBE8F8E8F749F89C" author="Jullien, J." box="[399,555,1796,1823]" pageId="13" pageNumber="336" pagination="180 - 183" refId="ref22649" refString="Jullien, J. (1890) Description d'un Bryozoaire nouveau du genre Rhabdopleura. Bulletin de la Societe Zoologique de France, 15, 180 - 183. https: // doi. org / 10.5962 / bhl. part. 18723" type="journal article" year="1890">Jullien (1890)</bibRefCitation>
and
<bibRefCitation id="EFB9DCAAFFC8B958E807F8E8F605F89C" author="Jullien, J. &amp; Calvet, L." box="[608,871,1796,1823]" pageId="13" pageNumber="336" pagination="1 - 188" refId="ref22692" refString="Jullien, J. &amp; Calvet, L. (1903) Bryozoaires provenant des campagnes de l'Hirondelle. Resultats des Campagnes scientifiques accompli par le Prince Albert I, 23, 1 - 188, 18 pls." type="journal article" year="1903">Jullien &amp; Calvet (1903)</bibRefCitation>
interpreted the thin brown lines as very fine tubes. These authors correctly illustrated an apparent gap (
<figureCitation id="1313BDDEFFC8B958E890F8C4F632F8C0" box="[759,848,1832,1859]" captionStart="FIGURE 4" captionStartId="10.[151,250,1795,1819]" captionTargetBox="[175,1413,182,1758]" captionTargetId="figure-28@10.[162,1426,181,1771]" captionTargetPageId="10" captionText="FIGURE 4. Original illustrations of described species of Rhabdopleura characterized by indirect inception of ringed erect tubes from creeping tubes via an intervening adnate portion that comprises a blind-ending branch. A, Rhabdopleura normani (from Allman 1869, pl. 8, fig. 1). Note that most of the colony is adherent; only the ringed tubes (rt) are erect, the proximal part of each side branch (ppsb) is adherent and the oblique sutures are not shown. B, schematic interpretative depiction of R. normani (after Andres 1977). C, D, Rhabdopleura grimaldii (from Jullien 1890, p. 181); note that ringed erect tubes (rt) are mostly lacking, with only the adherent proximal parts of side branches (ppsb) present. D is a cropped part of C to show the proximal parts of two side branches, with fusellar sutures and Julliens putative very fine tubes (vft)—actually the side walls of the tapering pectocaulus, the proximal end of which is transparent, hence the apparent gap between it and the pectocaulus of the creeping tube. E, F, Rhabdopleura striata (from Schepotieff 1909, pl. 7, figs 2, 7); E shows longitudinally striated erect parts of side branches; oblique sutures on prostrate parts omitted in E but shown in F, with characteristic web of fusellar sutures (wfs) in branch angles from main creeping tube. G, Rhabdopleura manubialis (from Jullien &amp; Calvet, 1903, pl. 1, fig. 2) with dormant bodies (db)." figureDoi="http://doi.org/10.5281/zenodo.10821369" httpUri="https://zenodo.org/record/10821369/files/figure.png" pageId="13" pageNumber="336">Fig. 4D</figureCitation>
) at the point where the brown pectocaulus diverges from the creeping tube into a side branch. At this point, the pectocaulus apparently becomes thinner, wider, and nearly transparent as it transitions into the side branch, with only its side walls clearly showing.
<bibRefCitation id="EFB9DCAAFFC8B958EEDEF89CF0FAF808" author="Schepotieff, A." box="[1209,1432,1904,1931]" pageId="13" pageNumber="336" pagination="463 - 534" refId="ref24624" refString="Schepotieff, A. (1907) Die Pterobranchier. Anatomische und histologische Untersuchungen uber Rhabdopleura normanii Allman und Cephalodiscus dodecalophus M'Int. 1. Teil. Rhabdopleura normanii Allman. Die Anatomie von Rhabdopleura. Zoologische Jahrbucher. Abteilung fur Anatomie und Ontogenie der Tiere, 23, 463 - 534, pls. 25 - 33. https: // doi. org / 10.5962 / bhl. part. 21044" type="journal article" year="1907">Schepotieffs (1907</bibRefCitation>
, pl. 33, figs 5, 9) cross sections of the pectocaulus in
<taxonomicName id="4C28DAD8FFC8B958E8BBF879F63BF82D" box="[732,857,1941,1966]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="13" pageNumber="336" phylum="Hemichordata" rank="species" species="normani">
<emphasis id="B95C7D49FFC8B958E8BBF879F63BF82D" box="[732,857,1941,1966]" italics="true" pageId="13" pageNumber="336">R. normani</emphasis>
</taxonomicName>
show side walls thickening at their base; this would account for the frontal appearance, in
<taxonomicName id="4C28DAD8FFC8B958E821F855F7ACF851" authorityName="Jullien" authorityYear="1890" box="[582,718,1976,2002]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="13" pageNumber="336" phylum="Hemichordata" rank="species" species="grimaldii">
<emphasis id="B95C7D49FFC8B958E821F855F7ACF851" box="[582,718,1976,2002]" italics="true" pageId="13" pageNumber="336">R. grimaldii</emphasis>
</taxonomicName>
, of the paired thin lines, each being a side wall. In this species, pectocaulus width prior to distal narrowing is 5174 (63) μm; the side walls (the so-called tubes très fins) are 1214 (13) μm (n = 4), too narrow for them to constitute converging paired pectocauli (an alternative interpretation). Presumably the distal tapering of the pectocaulus allows it to converge into the gymnocaulus of the zooid. [Actual zooids were not seen, however.] Erect ringed tubes are 92117 (104) μm diameter between fusellar collars (n = 8) and fusellus height is c. 1922 μm (
<emphasis id="B95C7D49FFCBB95BE84FFEEFF754FE9F" box="[552,566,259,284]" italics="true" pageId="14" pageNumber="337">n</emphasis>
= 2).
</paragraph>
<paragraph id="8B97A15BFFCBB95BEAA0FECBF1C2FE2E" blockId="14.[151,1437,151,465]" pageId="14" pageNumber="337">
Surface fusellar sutures of the adherent parts of side branches differ from those on primary creeping tubes, being closer together and having smaller zigzag angles. Two sequences of nine zigzag angles were measured, collectively ranging from 3360°, mean 46°, mode 35°. [In Julliens drawings, zigzag angles on side-branches are 4054°, mean 47°, mode 50° (
<emphasis id="B95C7D49FFCBB95BEB2EFE7FF435FE2F" box="[329,343,403,428]" italics="true" pageId="14" pageNumber="337">n</emphasis>
= 18), suggesting that his depictions of zigzag angles were pretty reliable.]
</paragraph>
<paragraph id="8B97A15BFFCBB95BEAA0FE5BF0E9FE52" blockId="14.[151,1437,151,465]" box="[199,1419,439,465]" pageId="14" pageNumber="337">
Our conclusion is that
<taxonomicName id="4C28DAD8FFCBB95BEBA2FE5BF72EFE52" authorityName="Jullien" authorityYear="1890" box="[453,588,439,465]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="14" pageNumber="337" phylum="Hemichordata" rank="species" species="grimaldii">
<emphasis id="B95C7D49FFCBB95BEBA2FE5BF72EFE52" box="[453,588,439,465]" italics="true" pageId="14" pageNumber="337">R. grimaldii</emphasis>
</taxonomicName>
is a highly distinctive species that cannot be synonymized with any other.
</paragraph>
</subSubSection>
</treatment>
</document>