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<mods:title id="226CEB01C90BC78BD6D31BAAFE204928">Systematic re-structure and new species of Sphaerodoridae (Annelida) after morphological revision and molecular phylogenetic analyses of the North East Atlantic fauna</mods:title>
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<mods:namePart id="545F08532A19299DBAEAB94F2E7648A4">Capa, Maria</mods:namePart>
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<mods:namePart id="71CDD45604796BE807C3E8FC5CFDF2A8">Bakken, Torkild</mods:namePart>
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<mods:namePart id="A9EC8CDAA3F0CC0FB3BE5BD375C0C41F">Moreira, Juan</mods:namePart>
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<treatment id="C734BBF54BFF9057AF453DA13CB0B57F" ID-GBIF-Taxon="157167804" LSID="urn:lsid:plazi:treatment:C734BBF54BFF9057AF453DA13CB0B57F" httpUri="http://treatment.plazi.org/id/C734BBF54BFF9057AF453DA13CB0B57F" lastPageId="23" lastPageNumber="24" pageId="21" pageNumber="22">
<subSubSection id="342823AA5975FF7580B43631517ACCAC" pageId="21" pageNumber="22" type="nomenclature">
<paragraph id="402947B12F326D05486C19E72E0F005C" pageId="21" pageNumber="22">
<taxonomicName id="DE4AC387455A776BE4423DEA364D821F" ID-CoL="XFW6" authority="Luetzen, 1961" authorityName="Luetzen" authorityYear="1961" class="Polychaeta" family="Sphaerodoridae" genus="Commensodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Commensodorum commensalis" order="Phyllodocida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="commensalis">
<pageBreakToken id="6C2FE202C4E667E84D9DA849DB9FCC63" pageId="21" pageNumber="22" start="start">Commensodorum</pageBreakToken>
commensalis (
<normalizedToken id="F3C761F6191E14D57CDC1C0BAA136956" originalValue="Lützen">Luetzen</normalizedToken>
, 1961)
</taxonomicName>
Figs 4E, F, 5C, 9
</paragraph>
</subSubSection>
<subSubSection id="DAB42F41BE0787A62E18A6E202621C5F" pageId="21" pageNumber="22" type="reference_group">
<paragraph id="23073E24973BF35642163A70BC653F8F" pageId="21" pageNumber="22">
<taxonomicName id="0464212F7AC123E6577329B7ACAD462D" class="Polychaeta" family="Sphaerodoridae" genus="Sphaerodoridium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphaerodoridium commensalis" order="Phyllodocida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="commensalis">Sphaerodoridium commensalis</taxonomicName>
<normalizedToken id="F7A0FAD7A08507083540220530A217F5" originalValue="Lützen">Luetzen</normalizedToken>
, 1961: 409-416, fig. 1.
</paragraph>
</subSubSection>
<subSubSection id="FB28D5E0BAF9D05921DF06855E9CCD0E" pageId="21" pageNumber="22" type="type locality">
<paragraph id="8CACEFE970FDA5A5E4A44ABC0553380B" pageId="21" pageNumber="22">Type locality.</paragraph>
<paragraph id="E25B3B36ED6507D247E8CFEC2E12DF35" pageId="21" pageNumber="22">
<normalizedToken id="31959D81DE3BF4223FAA9804EE0514B5" originalValue="Blåbergsholmen">Blabergsholmen</normalizedToken>
Island, Gullmarfjord, Sweden, 35 m.
</paragraph>
</subSubSection>
<subSubSection id="557308A775B848F03C1BAA63437F1824" pageId="21" pageNumber="22" type="materials_examined">
<paragraph id="FB49E7DF9AA7D9CAC4C0B049915B3986" pageId="21" pageNumber="22">Material examined.</paragraph>
<paragraph id="7BF7480AC6B019C3259BA9B3CFC75D73" pageId="21" pageNumber="22">
Holotype: ZMUC-POL-1984, Sweden: Gullmarfjord,
<normalizedToken id="700EA83A2986C2CFD0D4E0133E837D36" originalValue="Blåbergsholmen">Blabergsholmen</normalizedToken>
Island, 35 m, 30 Oct 1960.
</paragraph>
</subSubSection>
<subSubSection id="24430F83F84527E1EFA0F020468CA567" pageId="21" pageNumber="22" type="materials_examined">
<paragraph id="8BD3AB62907345FA66A01E569163B0E0" pageId="21" pageNumber="22">Additional material.</paragraph>
<paragraph id="68A257E45F119F02F8C8B19F4E4837A2" pageId="21" pageNumber="22">
(2 specs) Skagerrak, NTNU-VM 73780 (1 spec. on SEM stub) (1 spec.),
<normalizedToken id="35673EA240846C7F54D6C28FACD58336" originalValue="Lindön">Lindoen</normalizedToken>
,
<geoCoordinate id="414401F08A1785314BFE5D5C4A7AEB95" direction="north" orientation="latitude" precision="9" value="58.798332">58°47.90'N</geoCoordinate>
,
<geoCoordinate id="F324D12523B1C5CB9C55B5C9E5A637E7" direction="east" orientation="longitude" precision="9" value="11.158667">11°09.52'E</geoCoordinate>
, 46 m, 7 May 2008. ZMBN 127260 (1 spec.), Tvedestrand,
<geoCoordinate id="4DF825BDAA0F4196A87F673B7A83369B" direction="north" orientation="latitude" precision="1" value="58.565483">58°33.929'N</geoCoordinate>
,
<geoCoordinate id="10138A4124DA56B35F85DCD911CFE73F" direction="east" orientation="longitude" precision="1" value="9.003583">9°0.215'E</geoCoordinate>
, 34 m, 27 May 2011.
</paragraph>
</subSubSection>
<subSubSection id="B13ACD366BEB228D1D2A7A03E7F8FA95" pageId="21" pageNumber="22" type="diagnosis">
<paragraph id="E129F30D6D88AC16A48FE6CEBFD57856" pageId="21" pageNumber="22">Diagnosis.</paragraph>
<paragraph id="F2AC6DBB5CEB86EBA7E4BC428181680F" pageId="21" pageNumber="22">Body ellipsoid, up to 2.5 mm long. Head with short appendages, without spurs or basal papillae; antenniform papillae absent. Tubercles sessile, conical or pear-shaped, small, arranged in four longitudinal rows on dorsum, one transverse row per segment; except for first chaetiger with only two. Additional epithelial papillae minute over dorsal and ventral surfaces, in ca. 5-6 transverse rows per segment. Parapodia lacking papillae, acicular lobe, and ventral cirrus small and ellipsoid. Stout hooks in anterior chaetigers absent. All chaetae simple, unidentate, with broadened distal end, and serrated edge.</paragraph>
</subSubSection>
<subSubSection id="DF25B5744540272D50FD8D0743EC246B" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22" type="description">
<paragraph id="2DE9198EA42E42375DB1F204F0E2D213" pageId="21" pageNumber="22">Re-description of holotype.</paragraph>
<paragraph id="B5D6F4142622121929755ABE877FFB18" pageId="21" pageNumber="22">Measurements and general morphology. Holotype 2.5 mm long, 0.7 mm wide, with 17 chaetigers; body ellipsoid, with convex dorsum and flattened ventrum; segmentation slightly noticeable, especially on ventral side (Fig. 9A, F). Pigmentation absent on preserved material.</paragraph>
<caption id="2D291F34511E952389B6F81E19A006F6" pageId="21" pageNumber="22">
<paragraph id="A0A0ED98C7151AA1B5F3229A4D5E003D" pageId="21" pageNumber="22">
Figure 9.
<taxonomicName id="2AD426A208BEADB6B4A5DCC46B8358E0" class="Polychaeta" family="Sphaerodoridae" genus="Commensodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Commensodorum commensalis" order="Phyllodocida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="commensalis">Commensodorum commensalis</taxonomicName>
(Skagerrak, NTNU VM 73780). A Complete specimen, lateral view B anterior end, lateral view C mid-body chaetigers, lateral view D dorsal tubercle, mid-body chaetiger, detail E epithelium between dorsal tubercles, detail F parapodia, chaetigers 12-14, lateral view G parapodium, anterior chaetiger, lateral view H parapodium, mid-body chaetiger, anterior view I parapodium, posterior chaetiger J, K simple chaetae, anterior and posterior chaetigers L epithelium, detail, showing granules.
</paragraph>
</caption>
<paragraph id="024173FAB707DD8CB8BD2F06C58D399B" pageId="21" pageNumber="22">Head. Head fused to first chaetiger (Fig. 9A, B). Prostomial appendages conical, slightly longer than wide. A pair of palps, bigger than lateral and median antennae (Fig. 9B). Dorsal antenniform papillae absent. Few small hemispherical papillae scattered on head surface (only noticeable under SEM, Fig. 9B). Tentacular cirri ellipsoid, smaller than palps, similar in size and shape to median antenna (Fig. 9B). Eyes not observed.</paragraph>
<paragraph id="5BDBE9B1CAD18B350F306EC2E338357B" pageId="21" pageNumber="22">
Tubercles. Dorsum with four longitudinal rows of larger tubercles; in one transverse row per segment (Figs 5E, 9A, C). First chaetiger only with two tubercles. Distance between dorsalmost rows larger than there and lateral rows of tubercles. Tubercles sessile, ellipsoid or pear shaped (Fig. 9
<normalizedToken id="FA7E90B442B45443A061BC347ADB1BE9" originalValue="CE">C-E</normalizedToken>
), with some pores on distal end (Fig. 9D). Dorsal papillae minute, hemispherical, arranged in ca. six transverse rows per segment (Figs 4E, 9C, E). Ventrum with fewer and larger papillae, more abundant near parapodial bases, than in mid-ventral line; with no clear arrangement pattern (Figs 4F, 9F).
</paragraph>
<paragraph id="1403101D3163B7B8B78B9D2A57E1EECA" pageId="21" pageNumber="22">Parapodia. Parapodia conical, as long as wide in three or four anterior chaetigers, 2-3 times longer in medium and posterior chaetigers. Acicular lobes ellipsoid, from first chaetiger (Fig. 9G, H). Ventral cirri similar in shape and size as acicular lobe (Fig. 9H). Parapodial papillae absent (Fig. 5C).</paragraph>
<paragraph id="805373A36AD68F35198BE2DCB3973017" pageId="21" pageNumber="22">
Chaetae. All parapodia with 4-5 simple chaetae (Fig. 9
<normalizedToken id="74B83BF80CCCA7991084C1296465F0A2" originalValue="GI">G-I</normalizedToken>
); blade serrated on cutting edge and slight recurved distal tip; distal framed by thickened edges (Fig. 9
<normalizedToken id="464AE5305C786FADB0432A3107B97255" originalValue="IK">I-K</normalizedToken>
). One straight acicula per parapodium. Large, recurved hooks in the first chaetiger absent.
</paragraph>
<paragraph id="C3583BED2C6E45951F8613606EEEF764" pageId="21" pageNumber="22">Pygidium. Pygidium with two dorsolateral and one mid-ventral ellipsoid cirri, similar in size.</paragraph>
<paragraph id="0A116B45EB92D68BCD72B6CEA59A1396" pageId="22" pageNumber="23">
<pageBreakToken id="6AB3EAD25A4AFCB2DDA418E1EF12398A" pageId="22" pageNumber="23" start="start">Reproductive</pageBreakToken>
features. Holotype, gravid female, with oocytes measuring ca. 200
<normalizedToken id="2248A325E757B1357B40C8D1132D3229" originalValue="µm">µm</normalizedToken>
. Largest specimen also with oocytes, but smaller. Sexual structures or genital openings not observed.
</paragraph>
</subSubSection>
<subSubSection id="3DB564BE3E4070D15C5F7E70DE32C1C8" pageId="23" pageNumber="24" type="variation">
<paragraph id="0A172D3D4B45C2427B25E241F0E17FDE" pageId="23" pageNumber="24">
<pageBreakToken id="5DFABB73EA60598214C4DA49F8BF645B" pageId="23" pageNumber="24" start="start">Variation</pageBreakToken>
.
</paragraph>
<paragraph id="2C6C734C165175DB9B2F865045D3DA79" pageId="23" pageNumber="24">Largest specimens examined 4 mm long, 0.9 mm wide and 22 chaetigers. Smallest specimens with 19 chaetigers, 0.5 mm long. General morphology is homogenous among material studied. All specimens bear short prostomial appendages, ellipsoid or pear-shaped dorsal tubercles and minute epithelial papillae barely noticeable under stereomicroscope. One specimen (ZMUB 127260) is a gravid female with spheroid eggs occupying most of the body. Genital openings not observed in specimens examined.</paragraph>
</subSubSection>
<subSubSection id="8988C126B6AD373E681AA8F01C3E1259" pageId="23" pageNumber="24" type="remarks">
<paragraph id="DDCEC86F226D0538A9574C013DF4A8F6" pageId="23" pageNumber="24">Remarks.</paragraph>
<paragraph id="E9C7261BB30DE40E61F8916A917A5EAB" pageId="23" pageNumber="24">
Epithelium described as transparent in live specimens (
<bibRefCitation id="32E48806912288FE393791492E9ECD6E" author="Luetzen, J" journalOrPublisher="Cahiers de Biologie Marine" pageId="81" pageNumber="82" pagination="409 - 416" title="Sur une nouvelle espece de polychete Sphaerodoridiumcommensalis n. gen., n. spec. (PolychaetaErrantia, famille des Sphaerodoridae) vivant en commensal de Terebellidesstroemi Sars." volume="2" year="1961">
<normalizedToken id="EBBD0C06E4B7C8D8660B4ECC8C4A1755" originalValue="Lützen">Luetzen</normalizedToken>
1961
</bibRefCitation>
), allowing the observation of the nuchal organs (as pharyngeal glands) and the coiled gut. These are not visible in currently preserved specimens, with opaque epithelium. Original drawings of specimen with 22 chaetigers, does not correspond with the holotype. The body shape is ellipsoid, with blunt anterior and posterior ends, unlike the original description with tapering anterior end, probably due to contraction of specimens after fixation. Head appendages are short and ellipsoid in all specimens examined, not digitiform as in original description (
<bibRefCitation id="B2AF1A3723823EA2102166925A424F9D" author="Luetzen, J" journalOrPublisher="Cahiers de Biologie Marine" pageId="81" pageNumber="82" pagination="409 - 416" title="Sur une nouvelle espece de polychete Sphaerodoridiumcommensalis n. gen., n. spec. (PolychaetaErrantia, famille des Sphaerodoridae) vivant en commensal de Terebellidesstroemi Sars." volume="2" year="1961">
<normalizedToken id="6455405EB43BA6DB71AF720C59E6EEDB" originalValue="Lützen">Luetzen</normalizedToken>
1961
</bibRefCitation>
), also probably due to contraction. The main dorsal tubercles, four per chaetiger, were described as clavate ('forme de
<normalizedToken id="7EF997B787E9AA00A0F22A567D945904" originalValue="massue">massue'</normalizedToken>
), additionally other mid-dorsal
<normalizedToken id="9FD13DAA1EBA037E090686D1BA1C996E" originalValue="capsules">'capsules'</normalizedToken>
were described in anterior chaetigers, but they were not spotted in the material examined. Additional minute papillae are displayed forming ca. six transverse rows per segment.
</paragraph>
<paragraph id="E12E0B05C9EAEA799701CAFF3A21887E" pageId="23" pageNumber="24">
<taxonomicName id="739CA725CC20C4945B97E37649983F5C" class="Polychaeta" family="Sphaerodoridae" genus="Commensodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Commensodorum commensalis" order="Phyllodocida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="commensalis">Commensodorum commensalis</taxonomicName>
differs from other sphaerodorids in the presence of a unique combination of morphological features: four longitudinal rows of small dorsal tubercles, arranged in a single transverse row per segment, and the presence of simple chaetae. Sphaerodorids with simple chaetae comprise members of
<taxonomicName id="4C8993B9C123A0A542710FD9A10B6470" class="Polychaeta" family="Sphaerodoridae" genus="Euritmia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Euritmia" order="Phyllodocida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">Euritmia</taxonomicName>
, including the recently synonymised
<taxonomicName id="E1BDF7EB615F2D15868CD413D53ED928" class="Polychaeta" family="Sphaerodoridae" genus="Amacrodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amacrodorum" order="Phyllodocida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">Amacrodorum</taxonomicName>
(
<bibRefCitation id="280F06C18D27113CFD9DADAE2C46E32A" author="Capa, M" journalOrPublisher="ZooKeys" pageId="81" pageNumber="82" pagination="1 - 32" title="Sphaerodoridae (Annelida) of the deep Northwestern Atlantic, including remarkable new species of Euritmia and Sphaerephesia." url="https://doi.org/10.3897/zookeys.615.9530.figure1" volume="615" year="2016 b">Capa et al. 2016b</bibRefCitation>
), lacking large epithelial tubercles but with epithelium covered with small papillae. In addition, members of
<taxonomicName id="1B1CC5B7E49FCAC46C582A493DA60686" class="Polychaeta" family="Sphaerodoridae" genus="Sphaerodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphaerodorum" order="Phyllodocida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">Sphaerodorum</taxonomicName>
typically bear simple chaetae, but macrotubercles are arranged in two longitudinal rows and have clearly defined terminal papillae, absent in
<taxonomicName id="907158EC275B5BD5641CF3D0009FC2A4" class="Polychaeta" family="Sphaerodoridae" genus="Commensodorum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Commensodorum" order="Phyllodocida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">Commensodorum</taxonomicName>
, and bear in addition two longitudinal rows of dorsal microtubercles (small tubercles also provided with a terminal papilla).
</paragraph>
</subSubSection>
<subSubSection id="64FE618516DEBF9D8566D739CE6046BC" pageId="23" pageNumber="24" type="distribution">
<paragraph id="278948E98E282DAFDC123F06E242E072" pageId="23" pageNumber="24">Distribution.</paragraph>
<paragraph id="12CD6D441A1C9F5D89EEB7C12BCE3A6E" pageId="23" pageNumber="24">
Skagerrak,? United Kingdom (
<bibRefCitation id="7D48718249739F7ADACC76DE311025F3" author="Luetzen, J" journalOrPublisher="Cahiers de Biologie Marine" pageId="81" pageNumber="82" pagination="409 - 416" title="Sur une nouvelle espece de polychete Sphaerodoridiumcommensalis n. gen., n. spec. (PolychaetaErrantia, famille des Sphaerodoridae) vivant en commensal de Terebellidesstroemi Sars." volume="2" year="1961">
<normalizedToken id="E410253B4D610A588E0E39D479315E67" originalValue="Lützen">Luetzen</normalizedToken>
1961
</bibRefCitation>
,
<bibRefCitation id="59D8B8C11BAB594EB339C7E85491614A" author="Howson, CM" journalOrPublisher="Ulster Museum publication, no. 276. The Ulster Museum: Belfast" pageId="81" pageNumber="82" title="The species directory of the marine fauna and flora of the British Isles and surrounding seas." year="1997">Howson and Picton 1997</bibRefCitation>
,
<bibRefCitation id="5FE4F176EDF70D9E34B78883BC67F8C2" author="Frid, CLJ" journalOrPublisher="Journal of the Marine Biological Association of the United Kingdom" pageId="81" pageNumber="82" pagination="1 - 10" title="The North Sea benthic system: a 36 - year time-series." url="https://doi.org/10.1017/s0025315408002956" volume="89" year="2009">Frid et al. 2009</bibRefCitation>
, present study).
</paragraph>
</subSubSection>
<subSubSection id="E9E00A8ABEB02BE81CB2E6D45C472C58" pageId="23" pageNumber="24" type="habitat">
<paragraph id="A2FFD34A704B60B81744C345EBA14B01" pageId="23" pageNumber="24">Habitat.</paragraph>
<paragraph id="5D3CC746F4D42A227EB456B6D0B53496" pageId="23" pageNumber="24">
Originally described as commensal of
<taxonomicName id="A89BC70F8AE6C36640BFFE0E49BA7369" class="Polychaeta" family="Trichobranchidae" genus="Terebellides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Terebellides stroemii" order="Terebellida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="stroemii">Terebellides stroemii</taxonomicName>
Sars, 1835 (
<bibRefCitation id="0431327D94585453C7F1D69AEDC4667A" author="Luetzen, J" journalOrPublisher="Cahiers de Biologie Marine" pageId="81" pageNumber="82" pagination="409 - 416" title="Sur une nouvelle espece de polychete Sphaerodoridiumcommensalis n. gen., n. spec. (PolychaetaErrantia, famille des Sphaerodoridae) vivant en commensal de Terebellidesstroemi Sars." volume="2" year="1961">
<normalizedToken id="5EDF439639FEDF71FF0856192FDDCEAC" originalValue="Lützen">Luetzen</normalizedToken>
1961
</bibRefCitation>
), at 35 m. The species has only a few times been reported in the literature since and is not necessarily associated exclusively with
<taxonomicName id="362773277DBD1F86126C15A827CE71AB" class="Polychaeta" family="Trichobranchidae" genus="Terebellides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Terebellides" order="Terebellida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">Terebellides</taxonomicName>
(
<bibRefCitation id="25DECAFD20330506C1A94E4326264104" author="Frid, CLJ" journalOrPublisher="Journal of the Marine Biological Association of the United Kingdom" pageId="81" pageNumber="82" pagination="1 - 10" title="The North Sea benthic system: a 36 - year time-series." url="https://doi.org/10.1017/s0025315408002956" volume="89" year="2009">Frid et al. 2009</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>