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<mods:title id="AFDBA6CAC18F26EFCA7219D766D1147A">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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<mods:affiliation id="E15D8BDAEBB833B6D9F55F6BA34F40A3">National Institute of Water &amp; Atmospheric Research (NIWA), Private Bay 14901, Kilbirnie, Wellington 6241, New Zealand &amp; Department of Biological Sciences, National University of Singapore, 16 Science Drive 4, Singapore 117558, Singapore</mods:affiliation>
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<taxonomicName id="4C28DAD8FFCCB95CEAF0FCF2F4D5FCBA" ID-CoL="4RW6X" baseAuthorityName="Sars" baseAuthorityYear="1872" box="[151,439,798,825]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="9" pageNumber="332" phylum="Hemichordata" rank="species" species="mirabilis">
<emphasis id="B95C7D49FFCCB95CEAF0FCF2F4D5FCBA" bold="true" box="[151,439,798,825]" italics="true" pageId="9" pageNumber="332">Rhabdopleura mirabilis</emphasis>
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(M. Sars
<emphasis id="B95C7D49FFCCB95CE856FCF3F72BFCBA" bold="true" box="[561,585,799,825]" italics="true" pageId="9" pageNumber="332">in</emphasis>
G.O.
<bibRefCitation id="EFB9DCAAFFCCB95CE8F2FCF2F675FCBB" author="Sars, G. O." box="[661,791,798,824]" pageId="9" pageNumber="332" 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>
)
</paragraph>
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<subSubSection id="C332F2D0FFCCB95CE941FCF2F6F4FCBB" box="[806,918,798,824]" pageId="9" pageNumber="332" type="description">
<paragraph id="8B97A15BFFCCB95CE941FCF2F6F4FCBB" blockId="9.[151,918,798,825]" box="[806,918,798,824]" pageId="9" pageNumber="332">
(
<figureCitation id="1313BDDEFFCCB95CE948FCF2F6EEFCBB" box="[815,908,798,824]" captionStart="FIGURE 5" captionStartId="12.[151,250,1839,1863]" captionTargetBox="[201,1388,286,1803]" captionTargetId="figure-61@12.[186,1401,273,1816]" captionTargetPageId="12" captionText="FIGURE 5. Original illustrations of three described species of Rhabdopleura characterized by direct-frontal inception of ringed erect tubes (AC). Erect-tube inception in R. compacta (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. A, Rhabdopleura mirabilis (from Sars 1872, pl. 1, fig. 5). B, Rhabdopleura annulata (from Norman 1921, fig. 4). C, Rhabdopleura recondita (after Beli et al. 2018, fig. 2C). DF, Rhabdopleura compacta (respectively after Hincks 1880, pl. 72, fig. 8; Stebbing 1970b, fig. 1; Stebbing 1970a, fig. 3)." figureDoi="http://doi.org/10.5281/zenodo.10821371" httpUri="https://zenodo.org/record/10821371/files/figure.png" pageId="9" pageNumber="332">Fig. 5A</figureCitation>
)
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<paragraph id="8B97A15BFFCCB95CEAF0FC8BF06DFC02" blockId="9.[151,1437,870,2013]" box="[151,1295,870,897]" pageId="9" pageNumber="332">
<materialsCitation id="3B40AB06FFCCB95CEAF0FC8BF069FC02" box="[151,1291,870,897]" country="Norway" elevation="366" location="Skraaven" municipality="Lofoten Islands" pageId="9" pageNumber="332" specimenCount="1" typeStatus="holotype">
<emphasis id="B95C7D49FFCCB95CEAF0FC8BF454FC03" bold="true" box="[151,310,870,897]" pageId="9" pageNumber="332">
<typeStatus id="54931FF9FFCCB95CEAF0FC8BF5B0FC02" box="[151,210,871,897]" pageId="9" pageNumber="332">Type</typeStatus>
locality.
</emphasis>
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(= Skrova),
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,
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, at 100300 fm (c.
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) on mud
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.
</paragraph>
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<subSubSection id="C332F2D0FFCCB95CEAA0FC67F717FB6A" pageId="9" pageNumber="332" type="description">
<paragraph id="8B97A15BFFCCB95CEAA0FC67F717FB6A" blockId="9.[151,1437,870,2013]" pageId="9" pageNumber="332">
<emphasis id="B95C7D49FFCCB95CEAA0FC67F407FC27" bold="true" box="[199,357,906,933]" pageId="9" pageNumber="332">Key features.</emphasis>
Inception of ringed erect tubes is direct, with erect annulated zooid tubes produced directly from the creeping tube, not as blind branches. The creeping tube seldom bifurcates and is thickly covered by sediment particles, with erect tubes “always more or less bent in some part … sometimes like an S” (
<bibRefCitation id="EFB9DCAAFFCCB95CEEABFC3FF020FC6E" author="Sars, G. O." box="[1228,1346,979,1005]" pageId="9" pageNumber="332" 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>
, p. 26). Zigzag sutures were neither mentioned nor illustrated, presumably because of obscuring particles covering the creeping tube, or they may be lacking. The largest colony fragment obtained by Sars was
<quantity id="4CD00CBEFFCCB95CEE10FBF7F1CCFBB6" box="[1143,1198,1051,1077]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="9" pageNumber="332" unit="cm" value="4.0">4 cm</quantity>
long with erect tubes
<quantity id="4CD00CBEFFCCB95CEAF0FBD3F594FBDA" box="[151,246,1087,1113]" metricMagnitude="-3" metricUnit="m" metricValue="6.5" metricValueMax="7.0" metricValueMin="6.0" pageId="9" pageNumber="332" unit="mm" value="6.5" valueMax="7.0" valueMin="6.0">67 mm</quantity>
long and 200250 μm diameter. The living animal (arms, tentacles) was described as being covered by “intensely dark violet spots of colouring matter, which also occur on the buccal shield, and especially on its anterior freely projecting extremity, where they are very close together, forming a large, roundish dark spot” (
<emphasis id="B95C7D49FFCCB95CEF75FB6BF05CFB22" box="[1298,1342,1159,1185]" italics="true" pageId="9" pageNumber="332">ibid</emphasis>
., p. 34). Pigmentation thus differs from that in
<taxonomicName id="4C28DAD8FFCCB95CE826FB47F7DDFB47" ID-CoL="4RW6Y" box="[577,703,1195,1220]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="9" pageNumber="332" phylum="Hemichordata" rank="species" species="normani">
<emphasis id="B95C7D49FFCCB95CE826FB47F7DDFB47" box="[577,703,1195,1220]" italics="true" pageId="9" pageNumber="332">R. normani</emphasis>
</taxonomicName>
, which is black and orange-brown, conferring a “spotted leopard-like appearance” (
<bibRefCitation id="EFB9DCAAFFCCB95CEB06FB23F771FB6A" author="Lankester, E. R." box="[353,531,1231,1257]" pageId="9" pageNumber="332" pagination="622 - 647" refId="ref23121" refString="Lankester, E. R. (1884) A contribution to the knowledge of Rhabdopleura. Quarterly Journal of Microscopical Science, 24, 622 - 647, pls. 37 - 41. https: // doi. org / 10.1242 / jcs. s 2 - 24.96.622" type="journal article" year="1884">Lankester 1884</bibRefCitation>
, p. 630).
</paragraph>
</subSubSection>
<subSubSection id="C332F2D0FFCCB95CEAA0FB1EF58BF85E" pageId="9" pageNumber="332" type="discussion">
<paragraph id="8B97A15BFFCCB95CEAA0FB1EF714FA1E" blockId="9.[151,1437,870,2013]" pageId="9" pageNumber="332">
<emphasis id="B95C7D49FFCCB95CEAA0FB1EF426FA8F" bold="true" box="[199,324,1266,1292]" pageId="9" pageNumber="332">Comment.</emphasis>
Sars was well aware of Allmans species, concluding that it was “decidedly different” (
<bibRefCitation id="EFB9DCAAFFCCB95CEF43FB1FF0FAFA8E" author="Sars, G. O." box="[1316,1432,1267,1293]" pageId="9" pageNumber="332" 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>
, p. 27), lacking adherent particles and being “more strongly branched,” with shorter tubes and that “there appears some difference between the two species with regard to the manner of the division of the stem into chambers”;
<emphasis id="B95C7D49FFCCB95CEF1DFAD7F0FFFAD7" box="[1402,1437,1339,1364]" italics="true" pageId="9" pageNumber="332">i.e.</emphasis>
he noted the distinction between indirect-lateral and direct-frontal colony growth forms (as did
<bibRefCitation id="EFB9DCAAFFCCB95CEEBCFAB3F01BFAFA" author="Norman, J. R." box="[1243,1401,1375,1401]" pageId="9" pageNumber="332" 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>
, p. 99100), while not using that terminology.
</paragraph>
<paragraph id="8B97A15BFFCCB95CEAA0FA4BF7A3F93E" blockId="9.[151,1437,870,2013]" pageId="9" pageNumber="332">
Significantly, he also noted that, since the colony of
<taxonomicName id="4C28DAD8FFCCB95CE941FA4BF6CEFA42" baseAuthorityName="Sars" baseAuthorityYear="1872" box="[806,940,1447,1473]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="9" pageNumber="332" phylum="Hemichordata" rank="species" species="mirabilis">
<emphasis id="B95C7D49FFCCB95CE941FA4BF6CEFA42" box="[806,940,1447,1473]" italics="true" pageId="9" pageNumber="332">R. mirabilis</emphasis>
</taxonomicName>
is “very small and entirely colourless, it is rather difficult to discover. Its presence is, however, easily detected by stirring the washed mud in a fine sieve with a feather or other instrument, when irregular fibres will be noticed therein. These fibres, covered by particles of mud, Rhizopod-shells, and fragments of mussel shells, will prove to be the creeping stem, whereupon, by closer investigation, there will be discovered the small, transparent, perpendicularly projecting cells.” The muddy substratum is remarkable for an organism that is normally thought to require a firm surface for its creeping stem. Coupled with the differences in colony growth form and zooid pigmentation, our conclusion is that
<taxonomicName id="4C28DAD8FFCCB95CEF71F993F0F3F91A" authorityName="Sars" authorityYear="1872" box="[1302,1425,1663,1689]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="9" pageNumber="332" phylum="Hemichordata" rank="species" species="mirabili">
<emphasis id="B95C7D49FFCCB95CEF71F993F0F3F91A" box="[1302,1425,1663,1689]" italics="true" pageId="9" pageNumber="332">R. mirabili</emphasis>
</taxonomicName>
s cannot possibly be synonymous with
<taxonomicName id="4C28DAD8FFCCB95CE85BF94FF7D8F93F" box="[572,698,1699,1724]" class="Pterobranchia" family="Rhabdopleuridae" genus="Rhabdopleura" kingdom="Animalia" order="Rhabdopleurida" pageId="9" pageNumber="332" phylum="Hemichordata" rank="species" species="normani">
<emphasis id="B95C7D49FFCCB95CE85BF94FF7D8F93F" box="[572,698,1699,1724]" italics="true" pageId="9" pageNumber="332">R. normani</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B97A15BFFCCB95CEAA0F92BF58BF85E" blockId="9.[151,1437,870,2013]" pageId="9" pageNumber="332">
George Ossian Sars found the species in 1866. He later formally described it (
<bibRefCitation id="EFB9DCAAFFCCB95CEE41F92BF1F8F962" author="Sars, G. O." box="[1062,1178,1735,1761]" pageId="9" pageNumber="332" 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>
), stating that his father, Michael Sars, had noted it in his catalogue of deep-sea animals, compiled in 1868, under the denomination of
<taxonomicName id="4C28DAD8FFCCB95CEAFCF8E3F479F8AA" authorityName="Fabricius" authorityYear="1805" box="[155,283,1807,1833]" class="Amphibia" family="Ceratobatrachidae" genus="Halophilus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="332" phylum="Chordata" rank="genus">Halophilus</taxonomicName>
mirabilis.” This was in manuscript form as “Fortsatte Bemaerkninger om det dyriske Livs Udbredning i Havets Dybder” [= Continued Remarks on the Distribution of Animal Life in the Depths of the Sea]; it was not formally published and the combination
<taxonomicName id="4C28DAD8FFCCB95CE80BF8BBF635F8F2" box="[620,855,1879,1905]" class="Amphibia" family="Ceratobatrachidae" genus="Halophilus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="332" phylum="Chordata" rank="species" species="mirabilis">
<emphasis id="B95C7D49FFCCB95CE80BF8BBF635F8F2" box="[620,855,1879,1905]" italics="true" pageId="9" pageNumber="332">Halophilus mirabilis</emphasis>
</taxonomicName>
was a nomen nudum. Notwithstanding, G.O.
<bibRefCitation id="EFB9DCAAFFCCB95CEF0BF8BBF5B9F816" author="Sars, G. O." pageId="9" pageNumber="332" 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>
,
<bibRefCitation id="EFB9DCAAFFCCB95CEA8DF897F441F816" author="Sars, G. O." box="[234,291,1915,1941]" pageId="9" pageNumber="332" pagination="23 - 44" refId="ref24517" refString="Sars, G. O. (1874) On Rhabdopleura mirabilis (M. Sars). Quarterly Journal of Microscopical Science, New Series, 14, 23 - 44, pl. 1. https: // doi. org / 10.1242 / jcs. s 2 - 14.53.23" type="journal article" year="1874">1874</bibRefCitation>
) deliberately attributed authorship to his father. Accordingly, we consider formal authorship of
<taxonomicName id="4C28DAD8FFCCB95CEFE3F897F59EF83A" class="Amphibia" family="Ceratobatrachidae" genus="Halophilus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="332" phylum="Chordata" rank="species" species="mirabilis">
<emphasis id="B95C7D49FFCCB95CEFE3F897F59EF83A" italics="true" pageId="9" pageNumber="332">R. mirabilis</emphasis>
</taxonomicName>
to be M. Sars
<emphasis id="B95C7D49FFCCB95CEBCBF873F4A0F83B" box="[428,450,1951,1976]" italics="true" pageId="9" pageNumber="332">in</emphasis>
G.O.
<bibRefCitation id="EFB9DCAAFFCCB95CE86CF873F7E4F83A" author="Sars, G. O." box="[523,646,1951,1977]" pageId="9" pageNumber="332" 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>
. The account by G.O.
<bibRefCitation id="EFB9DCAAFFCCB95CE9E9F873F170F83A" author="Sars, G. O." box="[910,1042,1951,1977]" pageId="9" pageNumber="332" 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>
is repeated verbatim in G.O.
<bibRefCitation id="EFB9DCAAFFCCB95CEF0BF873F587F85E" author="Sars, G. O." pageId="9" pageNumber="332" pagination="23 - 44" refId="ref24517" refString="Sars, G. O. (1874) On Rhabdopleura mirabilis (M. Sars). Quarterly Journal of Microscopical Science, New Series, 14, 23 - 44, pl. 1. https: // doi. org / 10.1242 / jcs. s 2 - 14.53.23" type="journal article" year="1874">Sars (1874)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>