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<mods:title id="3C4BB7A4B85D79AFDB437E32C18A3C3E">Revision of the Laonice bahusiensis complex (Annelida: Spionidae) with a description of three new species</mods:title>
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<mods:affiliation id="F23584BAC6324DB72F7F81C67A7E03CB">University of Bologna, Department of Biological, Geological and Environmental Sciences, Via SantAlberto 163, 48123 Ravenna, Italy. barbara. mikac @ unibo. it; https: // orcid. org / 0000 - 0002 - 4516 - 0708</mods:affiliation>
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Sikorski, Radashevsky &amp; Nygren
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(
<figureCitation id="134B2A79FFF1FFFCFF5BF8BC3EE8F8BB" box="[159,250,1860,1887]" captionStart="FIGURE 5" captionStartId="12.[151,250,1000,1024]" captionTargetBox="[151,1436,417,977]" captionTargetId="figure-107@12.[151,1436,417,977]" captionTargetPageId="12" captionText="FIGURE 5. Maps showing records of Laonice bahusiensis complex species in the North East Atlantic Ocean (see Tables 1, S1S2 for details). A, Laonice bahusiensis: red star—type locality: Skagerrak, Gullmarfjord, Bohuslän, Sweden; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology. B, Laonice irinae n. sp.: red star—type locality: Norwegian Sea, Mikkelsøya, Norway; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology." figureDoi="http://doi.org/10.5281/zenodo.5069844" httpUri="https://zenodo.org/record/5069844/files/figure.png" pageId="11" pageNumber="264">Figs 5B</figureCitation>
,
<figureCitation id="134B2A79FFF1FFFCFEC2F8BC3F04F8BA" box="[262,278,1860,1886]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="11" pageNumber="264">6</figureCitation>
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<figureCitation id="134B2A79FFF1FFFCFEE6F8BD3F20F8BB" box="[290,306,1861,1887]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="11" pageNumber="264">7</figureCitation>
,
<figureCitation id="134B2A79FFF1FFFCFEFAF8BD3F5FF8BB" box="[318,333,1861,1887]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="11" pageNumber="264">8</figureCitation>
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<figureCitation id="134B2A79FFF1FFFCFE9EF8BD3F78F8BB" box="[346,362,1861,1887]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="11" pageNumber="264">9</figureCitation>
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<figureCitation id="134B2A79FFF1FFFCFEB2F8BC3FBBF8BB" box="[374,425,1860,1887]" captionStart="FIGURE 15" captionStartId="23.[151,250,1771,1795]" captionTargetBox="[151,1435,273,1748]" captionTargetId="figure-58@23.[151,1436,273,1748]" captionTargetPageId="23" captionText="FIGURE 15. Laonice spp. adult morphology.A, B, scheme showing the ratio length of prostomium in front of median eyes (l1) versus length of median eyes (l2): A, L. irinae n. sp., B, L. mediterranea n. sp. CH, L. mediterranea n. sp.: C, anterior end, dorsal view. D, chaetigers 2629, dorsal view, showing dorsal crests on postbranchiate region. E, F, parapodia and branchiae," figureDoi="http://doi.org/10.5281/zenodo.5069864" httpUri="https://zenodo.org/record/5069864/files/figure.png" pageId="11" pageNumber="264">15A</figureCitation>
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<figureCitation id="134B2A79FFF1FFFCFE71F8BD3FF4F8BB" box="[437,486,1861,1887]" captionStart="FIGURE 18" captionStartId="26.[151,250,1202,1226]" captionTargetBox="[151,1436,181,1178]" captionTargetId="figure-19@26.[151,1436,181,1178]" captionTargetPageId="26" captionText="FIGURE 18. Relationships between anterior position of lateral interneuropodial pouches (referring to the number of the chaetiger in front of the first pouch) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis. B, Laonice irinae n. sp. C, Laonice grimaldii n. sp. D, Laonice mediterranea n. sp." figureDoi="http://doi.org/10.5281/zenodo.5069870" httpUri="https://zenodo.org/record/5069870/files/figure.png" pageId="11" pageNumber="264">18B</figureCitation>
,
<figureCitation id="134B2A79FFF1FFFCFE35F8BD3C36F8BB" box="[497,548,1861,1887]" captionStart="FIGURE 19" captionStartId="26.[151,250,1807,1831]" captionTargetBox="[151,1436,1323,1784]" captionTargetId="figure-84@26.[151,1436,1323,1784]" captionTargetPageId="26" captionText="FIGURE 19. Relationships between the arithmetic difference between number of last branchiate chaetiger and length of nuchal organs (in chaetiger number) (Br-NO) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis (empty circles) and Laonice irinae n. sp. (filled circles). B, Laonice grimaldii n. sp. (empty circles) and Laonice mediterranea n. sp. (filled circles)." figureDoi="http://doi.org/10.5281/zenodo.5069872" httpUri="https://zenodo.org/record/5069872/files/figure.png" pageId="11" pageNumber="264">19A</figureCitation>
,
<tableCitation id="C6F20347FFF1FFFCFDEBF8BC3C90F8BB" box="[559,642,1860,1887]" captionStart="TABLE 3" captionStartId="25.[151,239,151,175]" captionText="TABLE 3. Determination coefficients (r2) between morphological and morphometric characters, width of chaetiger 7 (W7), and the distribution of branchiae (BR, referring to the number of the last branchiate chaetiger) in adults of Laonice spp. from European waters. For additional details see Figures 4, 8, 13, 1719. All correlations are significant (P &lt;0.05) except where indicated*." pageId="11" pageNumber="264">Table 3</tableCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C36A6577FFF1FFFCFF53F8753B20F80B" pageId="11" pageNumber="264" type="reference_group">
<paragraph id="8BCF36FCFFF1FFFCFF53F8753D7AF843" blockId="11.[151,1330,1932,2031]" box="[151,872,1932,1959]" pageId="11" pageNumber="264">
<taxonomicName id="4C704D7FFFF1FFFCFF53F8753C40F843" ID-CoL="6NX25" authority="Soderstrom, 1920" authorityName="Soderstrom" authorityYear="1920" box="[151,594,1932,1959]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="11" pageNumber="264" phylum="Annelida" rank="species" species="sarsi">
<emphasis id="B904EAEEFFF1FFFCFF53F8753F3CF842" box="[151,302,1933,1958]" italics="true" pageId="11" pageNumber="264">Laonice sarsi</emphasis>
<bibRefCitation id="EFE14B0DFFF1FFFCFEF2F8743FEFF843" author="Soderstrom, A." box="[310,509,1932,1959]" pageId="11" pageNumber="264" refId="ref18698" refString="Soderstrom, A. (1920) Studien uber die Polychatenfamilie Spionidae. Inaugural-Dissertation. Almquist &amp; Wicksells, Uppsala, 286 pp." type="book" year="1920">Söderström, 1920</bibRefCitation>
(
<emphasis id="B904EAEEFFF1FFFCFDC9F8753C2DF842" box="[525,575,1933,1958]" italics="true" pageId="11" pageNumber="264">Part</emphasis>
.)
</taxonomicName>
: 223225, figs 129, 130.
</paragraph>
<paragraph id="8BCF36FCFFF1FFFCFF53F8493A41F82F" blockId="11.[151,1330,1932,2031]" box="[151,1107,1968,1995]" pageId="11" pageNumber="264">
<treatmentCitationGroup id="AB6011D2FFF1FFFCFF53F8493A41F82F" box="[151,1107,1968,1995]" pageId="11" pageNumber="264">
<treatmentCitation id="0AD110EDFFF1FFFCFF53F8493C3FF82F" author="Ditlevsen, H." box="[151,557,1968,1995]" page="29" pageId="11" pageNumber="264" year="1929">
<taxonomicName id="4C704D7FFFF1FFFCFF53F8493C3FF82F" ID-CoL="3S88N" authority=": Ditlevsen 1929: 29" authorityName="Ditlevsen" authorityPageNumber="29" authorityYear="1929" box="[151,557,1968,1995]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="11" pageNumber="264" phylum="Annelida" rank="species" species="cirrata">
<emphasis id="B904EAEEFFF1FFFCFF53F8493F54F82E" box="[151,326,1969,1994]" italics="true" pageId="11" pageNumber="264">Laonice cirrata</emphasis>
:
<bibRefCitation id="EFE14B0DFFF1FFFCFE91F8483C3FF82F" author="Ditlevsen, H." box="[341,557,1968,1995]" pageId="11" pageNumber="264" pagination="1 - 83" refId="ref16802" refString="Ditlevsen, H. (1929) Polychaeta. Zoology of the Faroes, 16, 1 - 83." type="journal article" year="1929">Ditlevsen 1929: 29</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
<treatmentCitation id="0AD110EDFFF1FFFCFDFCF8483D31F82E" author="Kirkegaard, J. B." box="[568,803,1968,1994]" page="76" pageId="11" pageNumber="264" year="1969">
<bibRefCitation id="EFE14B0DFFF1FFFCFDFCF8483D31F82E" author="Kirkegaard, J. B." box="[568,803,1968,1994]" pageId="11" pageNumber="264" pagination="1 - 285" refId="ref17577" refString="Kirkegaard, J. B. (1969) A quantitative investigation of the central North Sea Polychaeta. Spolia Zoologica Musei Hauniensis, 29, 1 - 285." type="journal article" year="1969">Kirkegaard 1969: 76</bibRefCitation>
</treatmentCitation>
, fig. 40. Not M.
<bibRefCitation id="EFE14B0DFFF1FFFCFC19F8493A5DF82F" author="Sars, M." box="[989,1103,1968,1995]" pageId="11" pageNumber="264" pagination="121 - 211" refId="ref18123" refString="Sars, M. (1851) Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmarken. Nyt Magazin for Naturvidenskaberne, 6 (2), 121 - 211." type="journal article" year="1851">Sars 1851</bibRefCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BCF36FCFFF1FFFCFF53F82D3B20F80B" blockId="11.[151,1330,1932,2031]" box="[151,1330,2004,2031]" pageId="11" pageNumber="264">
<taxonomicName id="4C704D7FFFF1FFFCFF53F82D3C69F80B" ID-CoL="3S88G" authority=": Sikorski 2003" authorityName=": Sikorski" authorityYear="2003" box="[151,635,2004,2031]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="11" pageNumber="264" phylum="Annelida" rank="species" species="bahusiensis">
<emphasis id="B904EAEEFFF1FFFCFF53F82D3F69F80A" box="[151,379,2004,2030]" italics="true" pageId="11" pageNumber="264">Laonice bahusiensis</emphasis>
:
<bibRefCitation id="EFE14B0DFFF1FFFCFE4EF82C3C34F80B" author="Sikorski, A. V." box="[394,550,2004,2031]" pageId="11" pageNumber="264" pagination="316 - 345" refId="ref18276" refString="Sikorski, A. V. (2003) Laonice (Polychaeta, Spionidae) in the Arctic and the North Atlantic. Sarsia, 88 (5), 316 - 345. https: // doi. org / 10.1080 / 00364820310002551" type="journal article" year="2003">Sikorski 2003</bibRefCitation>
(
<emphasis id="B904EAEEFFF1FFFCFDF3F82D3C7BF80A" box="[567,617,2005,2030]" italics="true" pageId="11" pageNumber="264">Part</emphasis>
.)
</taxonomicName>
: 320325, figs 3AI, 4AB, 5AB,
<specimenCount id="9D76FD75FFF1FFFCFBD4F82C3A21F80A" box="[1040,1075,2004,2030]" pageId="11" pageNumber="264" type="generic">6F.</specimenCount>
Not
<bibRefCitation id="EFE14B0DFFF1FFFCFBAFF82C3B3CF80B" author="Soderstrom, A." box="[1131,1326,2004,2031]" pageId="11" pageNumber="264" refId="ref18698" refString="Soderstrom, A. (1920) Studien uber die Polychatenfamilie Spionidae. Inaugural-Dissertation. Almquist &amp; Wicksells, Uppsala, 286 pp." type="book" year="1920">Söderström 1920</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C36A6577FFF6FFF8FF53FF6F3AE8FEFA" lastPageId="15" lastPageNumber="268" pageId="12" pageNumber="265" type="materials_examined">
<paragraph id="8BCF36FCFFF6FFFBFF53FF6F3AACFF55" blockId="12.[151,1437,150,393]" box="[151,1214,150,177]" pageId="12" pageNumber="265">
<materialsCitation id="3B183CA1FFF6FFFBFF53FF6F3AACFF55" box="[151,1214,150,177]" country="Norway" elevation="38" latitude="68.6559" location="Norwegian Sea" longLatPrecision="5" longitude="14.7293" municipality="Mikkelsoya" pageId="12" pageNumber="265" specimenCount="38" typeStatus="holotype">
<emphasis id="B904EAEEFFF6FFFBFF53FF6F3F24FF54" bold="true" box="[151,310,150,177]" pageId="12" pageNumber="265">
<typeStatus id="54CB885EFFF6FFFBFF53FF6F3EC0FF55" box="[151,210,151,177]" pageId="12" pageNumber="265">Type</typeStatus>
locality.
</emphasis>
<collectingCountry id="F367766CFFF6FFFBFEF8FF6F3FBDFF55" box="[316,431,151,177]" name="Norway" pageId="12" pageNumber="265">NORWAY</collectingCountry>
,
<location id="8EAF6027FFF6FFFBFE79FF6F3C7EFF55" LSID="urn:lsid:plazi:treatment:03D987EAFFF1FFF8FF53F9043FCFF80A:8EAF6027FFF6FFFBFE79FF6F3C7EFF55" box="[445,620,151,177]" country="Norway" latitude="68.6559" longLatPrecision="5" longitude="14.7293" municipality="Mikkelsoya" name="Norwegian Sea" pageId="12" pageNumber="265">Norwegian Sea</location>
,
<collectingMunicipality id="6BABAC86FFF6FFFBFDB3FF6F3CECFF55" box="[631,766,151,177]" pageId="12" pageNumber="265">Mikkelsøya</collectingMunicipality>
, st. C2-1,
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,
<geoCoordinate id="EE44503BFFF6FFFBFC3DFF6F3A60FF55" box="[1017,1138,151,177]" degrees="14.7293" direction="east" orientation="longitude" pageId="12" pageNumber="265" precision="5" value="14.7293">14.7293°E</geoCoordinate>
,
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<quantity id="4C889B19FFF6FFFBFBBAFF6F3AA8FF55" box="[1150,1210,151,177]" metricMagnitude="1" metricUnit="m" metricValue="3.8" pageId="12" pageNumber="265" unit="m" value="38.0">
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</quantity>
.
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<paragraph id="8BCF36FCFFF6FFFBFF03FF433C3EFEF9" blockId="12.[151,1437,150,393]" pageId="12" pageNumber="265">
<emphasis id="B904EAEEFFF6FFFBFF03FF433F69FF30" bold="true" box="[199,379,186,213]" pageId="12" pageNumber="265">Type material.</emphasis>
<materialsCitation id="3B183CA1FFF6FFFBFE4DFF433FB3FF1D" collectionCode="ZMBN" pageId="12" pageNumber="265" specimenCode="ZMBN 135386" specimenCount="69" typeStatus="holotype">
<specimenCode id="DBD69E87FFF6FFFBFE4DFF433C29FF31" box="[393,571,186,213]" collectionCode="ZMBN" country="Norway" httpUri="http://grbio.org/cool/0tqf-efed" name="Museum of Zoology at the University of Bergen, Invertebrate Collection" pageId="12" pageNumber="265">ZMBN 135386</specimenCode>
(
<emphasis id="B904EAEEFFF6FFFBFD95FF433CAAFF31" bold="true" box="[593,696,187,213]" pageId="12" pageNumber="265">
<typeStatus id="54CB885EFFF6FFFBFD95FF433CAAFF31" box="[593,696,187,213]" pageId="12" pageNumber="265" type="holotype">holotype</typeStatus>
</emphasis>
), 116570, 135363135385, 135387135394, 135398, 136063 136067 (
<specimenCount id="9D76FD75FFF6FFFBFF3BFF273F8BFF1D" box="[255,409,223,249]" pageId="12" pageNumber="265" type="generic" typeStatus="paratypes">69 paratypes</specimenCount>
)
</materialsCitation>
;
<materialsCitation id="3B183CA1FFF6FFFBFE76FF273D4EFF1D" box="[434,860,223,249]" collectionCode="MIMB" pageId="12" pageNumber="265" specimenCode="MIMB 39043-39046" specimenCount="42" typeStatus="paratype">
<specimenCode id="DBD69E87FFF6FFFBFE76FF273CB7FF1D" box="[434,677,223,249]" collectionCode="MIMB" country="Russia" httpUri="http://grbio.org/cool/063z-3mpp" name="Museum of the Institute of Marine Biology" pageId="12" pageNumber="265">MIMB 3904339046</specimenCode>
(
<specimenCount id="9D76FD75FFF6FFFBFD7EFF273D41FF1D" box="[698,851,223,249]" pageId="12" pageNumber="265" type="generic" typeStatus="paratypes">42 paratypes</specimenCount>
)
</materialsCitation>
;
<materialsCitation id="3B183CA1FFF6FFFBFCA8FF273B8AFF1D" box="[876,1432,222,249]" collectionCode="NHMD" pageId="12" pageNumber="265" specimenCode="NHMD 108539, 108540, 108546" specimenCount="4" typeStatus="paratype">
<specimenCode id="DBD69E87FFF6FFFBFCA8FF273A37FF1D" box="[876,1061,222,249]" collectionCode="NHMD" pageId="12" pageNumber="265">NHMD 108539</specimenCode>
,
<specimenCode id="DBD69E87FFF6FFFBFBF1FF263A9EFF1D" box="[1077,1164,222,249]" collectionCode="NHMD" pageId="12" pageNumber="265">108540</specimenCode>
,
<specimenCode id="DBD69E87FFF6FFFBFB58FF263AE2FF1D" box="[1180,1264,222,249]" collectionCode="NHMD" pageId="12" pageNumber="265">108546</specimenCode>
(
<specimenCount id="9D76FD75FFF6FFFBFAC1FF273B82FF1D" box="[1285,1424,223,249]" pageId="12" pageNumber="265" type="generic" typeStatus="paratypes">4 paratypes</specimenCount>
)
</materialsCitation>
;
<materialsCitation id="3B183CA1FFF6FFFBFF53FEFB3C3BFEF9" box="[151,553,258,285]" collectionCode="UUZM" pageId="12" pageNumber="265" specimenCode="UUZM 52581, 52582" specimenCount="2" typeStatus="paratype">
<specimenCode id="DBD69E87FFF6FFFBFF53FEFB3F28FEF9" box="[151,314,258,285]" collectionCode="UUZM" country="Sweden" httpUri="http://grbio.org/cool/80z8-1h9s" name="Uppsala University, Zoological Museum" pageId="12" pageNumber="265">UUZM 52581</specimenCode>
,
<specimenCode id="DBD69E87FFF6FFFBFE81FEFA3F99FEF9" box="[325,395,258,285]" collectionCode="UUZM" country="Sweden" httpUri="http://grbio.org/cool/80z8-1h9s" name="Uppsala University, Zoological Museum" pageId="12" pageNumber="265">52582</specimenCode>
(
<specimenCount id="9D76FD75FFF6FFFBFE5EFEFB3C33FEF9" box="[410,545,259,285]" pageId="12" pageNumber="265" type="generic" typeStatus="paratypes">2 paratypes</specimenCount>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8BCF36FCFFF6FFFBFF03FEDE3FA8FE6D" blockId="12.[151,1437,150,393]" pageId="12" pageNumber="265">
<emphasis id="B904EAEEFFF6FFFBFF03FEDE3FB4FEA5" bold="true" box="[199,422,294,321]" pageId="12" pageNumber="265">Adult morphology.</emphasis>
Up to
<quantity id="4C889B19FFF6FFFBFE2AFEDE3C2EFEA5" box="[494,572,294,321]" metricMagnitude="-2" metricUnit="m" metricValue="4.5" pageId="12" pageNumber="265" unit="mm" value="45.0">45 mm</quantity>
long,
<quantity id="4C889B19FFF6FFFBFD44FEDE3CC7FEA5" box="[640,725,294,321]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="12" pageNumber="265" unit="mm" value="1.5">1.5 mm</quantity>
wide for 110 chaetigers; smallest examined complete individual
<quantity id="4C889B19FFF6FFFBFF53FEB33EF5FE81" box="[151,231,331,357]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="265" unit="mm" value="10.0">10 mm</quantity>
long,
<quantity id="4C889B19FFF6FFFBFEF4FEB33F95FE81" box="[304,391,331,357]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="12" pageNumber="265" unit="mm" value="0.3">0.3 mm</quantity>
wide for 60 chaetigers.
<typeStatus id="54CB885EFFF6FFFBFD5EFEB33D10FE81" box="[666,770,331,357]" pageId="12" pageNumber="265" type="holotype">Holotype</typeStatus>
<quantity id="4C889B19FFF6FFFBFCCFFEB33D49FE81" box="[779,859,331,357]" metricMagnitude="-2" metricUnit="m" metricValue="3.9" pageId="12" pageNumber="265" unit="mm" value="39.0">39 mm</quantity>
long,
<quantity id="4C889B19FFF6FFFBFC67FEB33DF7FE81" box="[931,997,331,357]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="265" unit="mm" value="1.0">1 mm</quantity>
wide for 102 chaetigers. Pigmentation absent on body and palps.
</paragraph>
<caption id="DF0F6674FFF6FFFBFF53FC103DACFB74" ID-DOI="http://doi.org/10.5281/zenodo.5069844" ID-Zenodo-Dep="5069844" httpUri="https://zenodo.org/record/5069844/files/figure.png" pageId="12" pageNumber="265" startId="12.[151,250,1000,1024]" targetBox="[151,1436,417,977]" targetPageId="12">
<paragraph id="8BCF36FCFFF6FFFBFF53FC103DACFB74" blockId="12.[151,1436,1000,1168]" pageId="12" pageNumber="265">
<emphasis id="B904EAEEFFF6FFFBFF53FC103F07FBE5" bold="true" box="[151,277,1000,1025]" pageId="12" pageNumber="265">FIGURE 5.</emphasis>
Maps showing records of
<taxonomicName id="4C704D7FFFF6FFFBFDEFFC113CE9FBE4" authorityName=": Sikorski" authorityYear="2003" box="[555,763,1000,1024]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="12" pageNumber="265" phylum="Annelida" rank="species" species="bahusiensis">
<emphasis id="B904EAEEFFF6FFFBFDEFFC113CE9FBE4" box="[555,763,1000,1024]" italics="true" pageId="12" pageNumber="265">Laonice bahusiensis</emphasis>
</taxonomicName>
complex species in the North East Atlantic Ocean (see Tables 1, S1S2 for details). A,
<taxonomicName id="4C704D7FFFF6FFFBFEB8FBF43C43FBC0" authorityName=": Sikorski" authorityYear="2003" box="[380,593,1036,1060]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="12" pageNumber="265" phylum="Annelida" rank="species" species="bahusiensis">
<emphasis id="B904EAEEFFF6FFFBFEB8FBF43C43FBC0" bold="true" box="[380,593,1036,1060]" italics="true" pageId="12" pageNumber="265">Laonice bahusiensis</emphasis>
</taxonomicName>
: red star—type locality: Skagerrak, Gullmarfjord, Bohuslän, Sweden; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology. B,
<emphasis id="B904EAEEFFF6FFFBFA99FBC93F2EFB89" bold="true" pageId="12" pageNumber="265">
<taxonomicName id="4C704D7FFFF6FFFBFA99FBC93EE8FB88" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="12" pageNumber="265" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF6FFFBFA99FBC93EE8FB88" bold="true" italics="true" pageId="12" pageNumber="265">Laonice irinae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2375795FFF6FFFBFEC6FBAD3F2EFB89" box="[258,316,1109,1133]" pageId="12" pageNumber="265" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
: red star—type locality: Norwegian Sea, Mikkelsøya, Norway; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology.
</paragraph>
</caption>
<paragraph id="8BCF36FCFFF6FFFBFF03FB473AF2FA1D" blockId="12.[151,1437,1215,2033]" pageId="12" pageNumber="265">
Prostomium approximately triangular, anteriorly wide, usually broadly rounded, occasionally truncate to slightly concave, fused with fronto-lateral margin of peristomium (
<figureCitation id="134B2A79FFF6FFFBFCDBFB1B3D8EFB19" box="[799,924,1251,1277]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Figs 6A, G</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFC62FB1B3DDEFB19" box="[934,972,1251,1277]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="12" pageNumber="265">7A</figureCitation>
), posteriorly extending over 36 chaetigers (to end of chaetiger
<quantity id="4C889B19FFF6FFFBFEBCFAFF3FA2FAC5" box="[376,432,1287,1313]" metricMagnitude="-1" metricUnit="m" metricValue="8.128" pageId="12" pageNumber="265" unit="in" value="32.0">32 in</quantity>
<typeStatus id="54CB885EFFF6FFFBFE73FAFF3C09FAC6" box="[439,539,1287,1314]" pageId="12" pageNumber="265" type="holotype">holotype</typeStatus>
) as a low narrow caruncle, shorter in small individuals (
<figureCitation id="134B2A79FFF6FFFBFB49FAFF3AF5FAC5" box="[1165,1255,1287,1313]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="12" pageNumber="265">Fig. 8A</figureCitation>
). Nuchal organs U-shaped ciliary bands on sides of caruncle (
<figureCitation id="134B2A79FFF6FFFBFD53FAD33CE4FAA1" box="[663,758,1323,1349]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Figs 6A</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFCC7FAD33D3AFAA1" box="[771,808,1323,1349]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="12" pageNumber="265">7A</figureCitation>
). Length of nuchal organs was strongly correlated with individual number of branchiate chaetigers (
<figureCitation id="134B2A79FFF6FFFBFD41FAB73CE3FA8D" box="[645,753,1359,1385]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="12" pageNumber="265">Fig. 8C, r</figureCitation>
<figureCitation id="134B2A79FFF6FFFBFD35FAB63CEBFAB8" box="[753,761,1358,1372]" captionStart="FIGURE 2" captionStartId="9.[151,250,1747,1771]" captionTargetBox="[152,1436,345,1724]" captionTargetId="figure-82@9.[151,1436,345,1724]" captionTargetPageId="9" captionText="FIGURE 2. Laonice bahusiensis adult morphology. A, anterior end, dorsal view. B, chaetigers 2733, dorsal view, showing posterior end of caruncle and nuchal organs, and dorsal crests on posterior branchiate and anterior abranchiate chaetigers. CE, parapodia and branchiae, left sides in frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining, ventral (F) and dorsal (G) view. A, B—ZMBN 135359; CE—ZMBN 135358; F, G—ZMBN 135358. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069828" httpUri="https://zenodo.org/record/5069828/files/figure.png" pageId="12" pageNumber="265">
<superScript id="7C059BB4FFF6FFFBFD35FAB63CEBFAB8" attach="left" box="[753,761,1358,1372]" fontSize="6" pageId="12" pageNumber="265">2</superScript>
</figureCitation>
= 0.8927, n = 57). Occipital antenna usually well developed (
<figureCitation id="134B2A79FFF6FFFBFF5BFA8B3EEFFA69" box="[159,253,1395,1421]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Figs 6A</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFECCFA8B3F3CFA69" box="[264,302,1395,1421]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="12" pageNumber="265">7A</figureCitation>
). Two pairs of red eyes (appearing almost black in formalin-fixed specimens) arranged trapezoidally, comprising one pair of large median eyes and one pair of small lateral eyes situated slightly anteriorly and set wider apart (often close to boundary between prostomium and peristomium and therefore not visible in dorsal view) (
<figureCitation id="134B2A79FFF6FFFBFAABFA433EAEFA1D" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Fig. 6A</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFF03FA263EEEFA1D" box="[199,252,1502,1529]" captionStart="FIGURE 15" captionStartId="23.[151,250,1771,1795]" captionTargetBox="[151,1435,273,1748]" captionTargetId="figure-58@23.[151,1436,273,1748]" captionTargetPageId="23" captionText="FIGURE 15. Laonice spp. adult morphology.A, B, scheme showing the ratio length of prostomium in front of median eyes (l1) versus length of median eyes (l2): A, L. irinae n. sp., B, L. mediterranea n. sp. CH, L. mediterranea n. sp.: C, anterior end, dorsal view. D, chaetigers 2629, dorsal view, showing dorsal crests on postbranchiate region. E, F, parapodia and branchiae," figureDoi="http://doi.org/10.5281/zenodo.5069864" httpUri="https://zenodo.org/record/5069864/files/figure.png" pageId="12" pageNumber="265">15A</figureCitation>
). Palps as long as 512 chaetigers, with deep frontal longitudinal groove lined with cilia.
</paragraph>
<paragraph id="8BCF36FCFFF6FFFBFF03F9FB3CB2F911" blockId="12.[151,1437,1215,2033]" pageId="12" pageNumber="265">
Chaetiger 1 with well-developed capillary chaetae and small postchaetal lamellae in both rami; notopodial lamellae triangular; neuropodial lamellae rounded.All notopodia with capillary chaetae only. Low prechaetal lamellae present in noto- and neuropodia on anterior chaetigers after chaetiger 1. Notopodial postchaetal lamellae large, leaf-like on branchiate chaetigers, usually largest on chaetiger 4, not overlapping middorsally, greatly diminishing in size on posterior abranchiate chaetigers; lamellae on at least ten anterior chaetigers with terminal acute peaks on upper tips (
<figureCitation id="134B2A79FFF6FFFBFED2F94F3FC0F935" box="[278,466,1719,1745]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Fig. 6A, CE, G</figureCitation>
). Neuropodial postchaetal lamellae ear-like on branchiate chaetigers, greatly diminishing in size on posterior abranchiate chaetigers.
</paragraph>
<paragraph id="8BCF36FCFFF6FFFBFF03F9073FBAF829" blockId="12.[151,1437,1215,2033]" pageId="12" pageNumber="265">
Branchiae from chaetiger 2, up to 32 pairs (on chaetigers
<quantity id="4C889B19FFF6FFFBFC84F9073D86F8FD" box="[832,916,1791,1817]" metricMagnitude="-1" metricUnit="m" metricValue="4.064" metricValueMax="7.62" metricValueMin="0.508" pageId="12" pageNumber="265" unit="in" value="16.0" valueMax="30.0" valueMin="2.0">230 in</quantity>
<typeStatus id="54CB885EFFF6FFFBFC5DF9073A12F8FE" box="[921,1024,1791,1818]" pageId="12" pageNumber="265" type="holotype">holotype</typeStatus>
); first pair shorter or similar in length to notopodial postchaetal lamellae of chaetiger 2; from chaetigers 56 branchiae longer (up to twice as long) than notopodial postchaetal lamellae (
<figureCitation id="134B2A79FFF6FFFBFDCBF8BF3C9EF885" box="[527,652,1863,1889]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Fig. 6E, G</figureCitation>
), gradually diminishing in size on succeeding chaetigers, posteriorly disappearing 06 (usually 13) chaetigers before end of caruncle and nuchal organs, or one chaetiger after end of nuchal organs (
<figureCitation id="134B2A79FFF6FFFBFE81F8773FB0F84D" box="[325,418,1935,1961]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="12" pageNumber="265">Figs 6B</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFE6BF8773FC0F84D" box="[431,466,1935,1961]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="12" pageNumber="265">7B</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFE1BF8773C34F84D" box="[479,550,1935,1961]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="12" pageNumber="265">8A, C</figureCitation>
,
<figureCitation id="134B2A79FFF6FFFBFDF7F8773C75F84D" box="[563,615,1935,1961]" captionStart="FIGURE 19" captionStartId="26.[151,250,1807,1831]" captionTargetBox="[151,1436,1323,1784]" captionTargetId="figure-84@26.[151,1436,1323,1784]" captionTargetPageId="26" captionText="FIGURE 19. Relationships between the arithmetic difference between number of last branchiate chaetiger and length of nuchal organs (in chaetiger number) (Br-NO) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis (empty circles) and Laonice irinae n. sp. (filled circles). B, Laonice grimaldii n. sp. (empty circles) and Laonice mediterranea n. sp. (filled circles)." figureDoi="http://doi.org/10.5281/zenodo.5069872" httpUri="https://zenodo.org/record/5069872/files/figure.png" pageId="12" pageNumber="265">19A</figureCitation>
). Individual number of branchiae was strongly correlated with length of nuchal organs (
<figureCitation id="134B2A79FFF6FFFBFE86F84B3F89F829" box="[322,411,1971,1997]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="12" pageNumber="265">Fig. 8C</figureCitation>
).
</paragraph>
<paragraph id="8BCF36FCFFF6FFFAFF03F82F3A85FF1E" blockId="12.[151,1437,1215,2033]" lastBlockId="13.[151,1436,151,357]" lastPageId="13" lastPageNumber="266" pageId="12" pageNumber="265">
Dorsal transverse crests one per chaetiger, beginning 14 chaetigers before, and arranged on up to 15 chaetigers after the posterior end of nuchal organs (
<figureCitation id="134B2A79FFF7FFFAFDA2FF6F3CD0FF55" box="[614,706,151,177]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="13" pageNumber="266">Figs 6B</figureCitation>
,
<figureCitation id="134B2A79FFF7FFFAFD0BFF6F3CE6FF55" box="[719,756,151,177]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="13" pageNumber="266">7B</figureCitation>
). Several anterior dorsal crests interrupted by nuchal organs (nuchal organs sometimes are interrupted by the dorsal crests on one or two most posterior chaetigers); on succeeding chaetigers crests continuous, interconnecting notopodial postchaetal lamellae (
<figureCitation id="134B2A79FFF7FFFAFBF5FF273A98FF1E" box="[1073,1162,223,250]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="13" pageNumber="266">Fig. 6B</figureCitation>
).
</paragraph>
<paragraph id="8BCF36FCFFF7FFFAFF03FEFB3EFDFE81" blockId="13.[151,1436,151,357]" pageId="13" pageNumber="266">
Lateral interneuropodial pouches from chaetigers 620 (from chaetiger
<quantity id="4C889B19FFF7FFFAFC27FEFB3A0EFEF9" box="[995,1052,259,285]" metricMagnitude="-1" metricUnit="m" metricValue="3.302" pageId="13" pageNumber="266" unit="in" value="13.0">13 in</quantity>
<typeStatus id="54CB885EFFF7FFFAFBE6FEFB3A94FEFA" box="[1058,1158,259,286]" pageId="13" pageNumber="266" type="holotype">holotype</typeStatus>
) to end of body. Anterior start of pouches was moderately correlated with individual number of branchiate chaetigers (
<figureCitation id="134B2A79FFF7FFFAFB63FEDF3B30FEA5" box="[1191,1314,295,321]" captionStart="FIGURE 18" captionStartId="26.[151,250,1202,1226]" captionTargetBox="[151,1436,181,1178]" captionTargetId="figure-19@26.[151,1436,181,1178]" captionTargetPageId="26" captionText="FIGURE 18. Relationships between anterior position of lateral interneuropodial pouches (referring to the number of the chaetiger in front of the first pouch) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis. B, Laonice irinae n. sp. C, Laonice grimaldii n. sp. D, Laonice mediterranea n. sp." figureDoi="http://doi.org/10.5281/zenodo.5069870" httpUri="https://zenodo.org/record/5069870/files/figure.png" pageId="13" pageNumber="266">Fig. 18B, r</figureCitation>
<figureCitation id="134B2A79FFF7FFFAFAE6FEDE3B38FED0" box="[1314,1322,294,308]" captionStart="FIGURE 2" captionStartId="9.[151,250,1747,1771]" captionTargetBox="[152,1436,345,1724]" captionTargetId="figure-82@9.[151,1436,345,1724]" captionTargetPageId="9" captionText="FIGURE 2. Laonice bahusiensis adult morphology. A, anterior end, dorsal view. B, chaetigers 2733, dorsal view, showing posterior end of caruncle and nuchal organs, and dorsal crests on posterior branchiate and anterior abranchiate chaetigers. CE, parapodia and branchiae, left sides in frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining, ventral (F) and dorsal (G) view. A, B—ZMBN 135359; CE—ZMBN 135358; F, G—ZMBN 135358. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069828" httpUri="https://zenodo.org/record/5069828/files/figure.png" pageId="13" pageNumber="266">
<superScript id="7C059BB4FFF7FFFAFAE6FEDE3B38FED0" attach="left" box="[1314,1322,294,308]" fontSize="6" pageId="13" pageNumber="266">2</superScript>
</figureCitation>
= 0.6665, n = 57).
</paragraph>
<caption id="DF0F6674FFF7FFFAFF53F8C13AB7F85A" ID-DOI="http://doi.org/10.5281/zenodo.5069846" ID-Zenodo-Dep="5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="13" pageNumber="266" startId="13.[151,250,1849,1873]" targetBox="[154,1436,381,1826]" targetPageId="13">
<paragraph id="8BCF36FCFFF7FFFAFF53F8C13AB7F85A" blockId="13.[151,1437,1849,1982]" pageId="13" pageNumber="266">
<emphasis id="B904EAEEFFF7FFFAFF53F8C13FE7F8B6" bold="true" box="[151,501,1849,1874]" pageId="13" pageNumber="266">
FIGURE 6.
<taxonomicName id="4C704D7FFFF7FFFAFEDFF8C23FA6F8B6" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[283,436,1850,1874]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="13" pageNumber="266" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF7FFFAFEDFF8C23FA6F8B6" bold="true" box="[283,436,1850,1874]" italics="true" pageId="13" pageNumber="266">Laonice irinae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2375795FFF7FFFAFE7FF8C23FE7F8B6" box="[443,501,1850,1874]" pageId="13" pageNumber="266" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view. A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm.
</paragraph>
</caption>
<paragraph id="8BCF36FCFFF4FFF9FF03FF6F3FF5FEF9" blockId="14.[151,1437,151,573]" pageId="14" pageNumber="267">
Sabre chaetae in neuropodia from chaetigers 921 (from chaetiger
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<typeStatus id="54CB885EFFF4FFF9FBC0FF6F3A78FF56" box="[1028,1130,151,178]" pageId="14" pageNumber="267" type="holotype">holotype</typeStatus>
), from more anterior chaetigers in small individuals (
<figureCitation id="134B2A79FFF4FFF9FE0AFF433C34FF31" box="[462,550,187,213]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="14" pageNumber="267">Fig. 8B</figureCitation>
),
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a tuft; chaetae with weak granulation on shaft, up to twice as long as hooks. Anterior start of sabre chaetae was moderately correlated with individual number of branchiate chaetigers (
<figureCitation id="134B2A79FFF4FFF9FF64FEFB3F1FFEF9" box="[160,269,259,285]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="14" pageNumber="267">Fig. 8D, r</figureCitation>
<figureCitation id="134B2A79FFF4FFF9FEC9FEFA3F07FEF4" box="[269,277,258,272]" captionStart="FIGURE 2" captionStartId="9.[151,250,1747,1771]" captionTargetBox="[152,1436,345,1724]" captionTargetId="figure-82@9.[151,1436,345,1724]" captionTargetPageId="9" captionText="FIGURE 2. Laonice bahusiensis adult morphology. A, anterior end, dorsal view. B, chaetigers 2733, dorsal view, showing posterior end of caruncle and nuchal organs, and dorsal crests on posterior branchiate and anterior abranchiate chaetigers. CE, parapodia and branchiae, left sides in frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining, ventral (F) and dorsal (G) view. A, B—ZMBN 135359; CE—ZMBN 135358; F, G—ZMBN 135358. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069828" httpUri="https://zenodo.org/record/5069828/files/figure.png" pageId="14" pageNumber="267">
<superScript id="7C059BB4FFF4FFF9FEC9FEFA3F07FEF4" attach="left" box="[269,277,258,272]" fontSize="6" pageId="14" pageNumber="267">2</superScript>
</figureCitation>
= 0.5057, n = 58).
</paragraph>
<paragraph id="8BCF36FCFFF4FFF9FF03FEDF3C4DFDD9" blockId="14.[151,1437,151,573]" pageId="14" pageNumber="267">
Hooded hooks in neuropodia from chaetigers 1435 (from chaetiger
<quantity id="4C889B19FFF4FFF9FC0EFEDF3A11FEA5" box="[970,1027,295,321]" metricMagnitude="-1" metricUnit="m" metricValue="7.366" pageId="14" pageNumber="267" unit="in" value="29.0">29 in</quantity>
<typeStatus id="54CB885EFFF4FFF9FBCEFEDF3A62FEA6" box="[1034,1136,295,322]" pageId="14" pageNumber="267" type="holotype">holotype</typeStatus>
), from more anterior chaetigers in small individuals (
<figureCitation id="134B2A79FFF4FFF9FE16FEB33C3EFE81" box="[466,556,331,357]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="14" pageNumber="267">Fig. 8B</figureCitation>
), up to
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a series, accompanied by inferior sabre chaetae and alternating capillaries throughout body. Alternating capillaries with very narrow limbation, up to 1.5 times as long as hooks, up to
<quantity id="4C889B19FFF4FFF9FF1DFE6B3F01FE49" box="[217,275,403,429]" metricMagnitude="-1" metricUnit="m" metricValue="2.54" pageId="14" pageNumber="267" unit="in" value="10.0">10 in</quantity>
a series, in middle and posterior chaetigers situated in upper part of hook row. Hooks quinquedentate, with two pairs of upper teeth arranged in two vertical rows above main fang (
<figureCitation id="134B2A79FFF4FFF9FBC2FE4F3A4DFE35" box="[1030,1119,439,465]" captionStart="FIGURE 7" captionStartId="14.[151,250,1114,1138]" captionTargetBox="[162,1424,598,1091]" captionTargetId="figure-222@14.[162,1424,597,1091]" captionTargetPageId="14" captionText="FIGURE 7. Laonice irinae n. sp. adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open.Abbreviations: an—occipital antenna; br—branchia; ca—caruncle; lp—lateral interneuropodial pouch; no—nototrochs; nu—doublerow nuchal organ; pe—peristomium; pr—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm." figureDoi="http://doi.org/10.5281/zenodo.5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="14" pageNumber="267">Fig. 7C</figureCitation>
), or quadridentate, with one pair of upper teeth and a single superior median tooth above main fang; shaft slightly curved, slightly narrowed at level of body cuticle (
<figureCitation id="134B2A79FFF4FFF9FE4EFE073FCCFDFD" box="[394,478,511,537]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9F</figureCitation>
). Anterior start of hooks was strongly correlated with individual number of branchiate chaetigers (
<figureCitation id="134B2A79FFF4FFF9FEDCFDDB3F94FDD9" box="[280,390,547,573]" captionStart="FIGURE 8" captionStartId="15.[151,250,1465,1489]" captionTargetBox="[177,1411,489,1442]" captionTargetId="figure-176@15.[177,1411,489,1442]" captionTargetPageId="15" captionText="FIGURE 8. Laonice irinae n. sp. adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles)." figureDoi="http://doi.org/10.5281/zenodo.5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="14" pageNumber="267">Fig. 8D, r</figureCitation>
<figureCitation id="134B2A79FFF4FFF9FE42FDDA3F9CFDD4" box="[390,398,546,560]" captionStart="FIGURE 2" captionStartId="9.[151,250,1747,1771]" captionTargetBox="[152,1436,345,1724]" captionTargetId="figure-82@9.[151,1436,345,1724]" captionTargetPageId="9" captionText="FIGURE 2. Laonice bahusiensis adult morphology. A, anterior end, dorsal view. B, chaetigers 2733, dorsal view, showing posterior end of caruncle and nuchal organs, and dorsal crests on posterior branchiate and anterior abranchiate chaetigers. CE, parapodia and branchiae, left sides in frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining, ventral (F) and dorsal (G) view. A, B—ZMBN 135359; CE—ZMBN 135358; F, G—ZMBN 135358. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069828" httpUri="https://zenodo.org/record/5069828/files/figure.png" pageId="14" pageNumber="267">
<superScript id="7C059BB4FFF4FFF9FE42FDDA3F9CFDD4" attach="left" box="[390,398,546,560]" fontSize="6" pageId="14" pageNumber="267">2</superScript>
</figureCitation>
= 0.8138, n = 58).
</paragraph>
<caption id="DF0F6674FFF4FFF9FF53FBA23B01FB3B" ID-DOI="http://doi.org/10.5281/zenodo.5069848" ID-Zenodo-Dep="5069848" httpUri="https://zenodo.org/record/5069848/files/figure.png" pageId="14" pageNumber="267" startId="14.[151,250,1114,1138]" targetBox="[162,1424,598,1091]" targetPageId="14">
<paragraph id="8BCF36FCFFF4FFF9FF53FBA23B01FB3B" blockId="14.[151,1437,1114,1247]" pageId="14" pageNumber="267">
<emphasis id="B904EAEEFFF4FFF9FF53FBA23FE0FB97" bold="true" box="[151,498,1114,1139]" pageId="14" pageNumber="267">
FIGURE 7.
<taxonomicName id="4C704D7FFFF4FFF9FEDEFBA33FA0FB97" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[282,434,1115,1139]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="14" pageNumber="267" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF4FFF9FEDEFBA33FA0FB97" bold="true" box="[282,434,1115,1139]" italics="true" pageId="14" pageNumber="267">Laonice irinae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2375795FFF4FFF9FE7DFBA33FE0FB97" box="[441,498,1115,1139]" pageId="14" pageNumber="267" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
adult morphology (SEM). A, anterior end, dorsal view. B, midbody chaetigers, dorso-lateral view, showing posterior end of caruncle with nuchal organs (arrow). C, distal part of neuropodial hook with hood open. Abbreviations:
<emphasis id="B904EAEEFFF4FFF9FF30FB5B3F1FFB5E" box="[244,269,1187,1210]" italics="true" pageId="14" pageNumber="267">an</emphasis>
—occipital antenna;
<emphasis id="B904EAEEFFF4FFF9FE24FB5B3FE5FB5F" box="[480,503,1187,1211]" italics="true" pageId="14" pageNumber="267">br</emphasis>
—branchia;
<emphasis id="B904EAEEFFF4FFF9FDB0FB5B3C9EFB5E" box="[628,652,1187,1210]" italics="true" pageId="14" pageNumber="267">ca</emphasis>
—caruncle;
<emphasis id="B904EAEEFFF4FFF9FCC3FB5B3D09FB5F" box="[775,795,1187,1211]" italics="true" pageId="14" pageNumber="267">lp</emphasis>
—lateral interneuropodial pouch;
<emphasis id="B904EAEEFFF4FFF9FBB6FB5B3A99FB5E" box="[1138,1163,1187,1210]" italics="true" pageId="14" pageNumber="267">no</emphasis>
—nototrochs;
<emphasis id="B904EAEEFFF4FFF9FAD8FB5B3B27FB5E" box="[1308,1333,1187,1210]" italics="true" pageId="14" pageNumber="267">nu</emphasis>
—doublerow nuchal organ;
<emphasis id="B904EAEEFFF4FFF9FE91FB3F3F7FFB3A" box="[341,365,1223,1246]" italics="true" pageId="14" pageNumber="267">pe</emphasis>
—peristomium;
<emphasis id="B904EAEEFFF4FFF9FDD0FB3F3C39FB3A" box="[532,555,1223,1246]" italics="true" pageId="14" pageNumber="267">pr</emphasis>
—prostomium. AC—VIR 23519. Scale bars: A, B = 200 µm; C = 2 µm.
</paragraph>
</caption>
<paragraph id="8BCF36FCFFF4FFF9FF03FAF53B46FAAF" blockId="14.[151,1437,1293,2039]" pageId="14" pageNumber="267">
Pygidium with up to eleven pairs of cirri (seven pairs in
<typeStatus id="54CB885EFFF4FFF9FC82FAF53DB8FACC" box="[838,938,1293,1320]" pageId="14" pageNumber="267" type="holotype">holotype</typeStatus>
) arranged around terminal anus, comprising one pair of ventral cirri and up to ten pairs of thinner and longer dorsal cirri; fewer cirri in small individuals.
</paragraph>
<paragraph id="8BCF36FCFFF4FFF9FF03FAAD3CD7FA8B" blockId="14.[151,1437,1293,2039]" box="[199,709,1365,1391]" pageId="14" pageNumber="267">Digestive tract without gizzard-like structure.</paragraph>
<paragraph id="8BCF36FCFFF4FFF9FF03FA813FD1FA53" blockId="14.[151,1437,1293,2039]" pageId="14" pageNumber="267">Nephridia from chaetiger 4 to chaetigers 2124, present in all anterior sterile chaetigers except chaetigers 13, fewer in small individuals.</paragraph>
<paragraph id="8BCF36FCFFF4FFF9FF03FA393B0FF8A7" blockId="14.[151,1437,1293,2039]" pageId="14" pageNumber="267">
<emphasis id="B904EAEEFFF4FFF9FF03FA393F61FA3F" bold="true" box="[199,371,1473,1499]" pageId="14" pageNumber="267">Reproduction.</emphasis>
<taxonomicName id="4C704D7FFFF4FFF9FEBDFA3A3C0FFA3F" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[377,541,1474,1499]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="14" pageNumber="267" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF4FFF9FEBDFA3A3C0FFA3F" box="[377,541,1474,1499]" italics="true" pageId="14" pageNumber="267">Laonice irinae</emphasis>
</taxonomicName>
<emphasis id="B904EAEEFFF4FFF9FDE7FA393C70FA3F" bold="true" box="[547,610,1473,1499]" pageId="14" pageNumber="267">
<taxonomicNameLabel id="A2375795FFF4FFF9FDE7FA393C70FA3F" box="[547,610,1473,1499]" pageId="14" pageNumber="267" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
is dioecious. Gametes develop in both females and males from chaetigers 2325 through most of the body. Oogenesis is entirely intraovarian; vitellogenesis occurs when oocytes grow while attached to segmental blood vessels (
<figureCitation id="134B2A79FFF4FFF9FDFFF9F13C87F9C7" box="[571,661,1545,1571]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9B</figureCitation>
). Developed oocytes are released into the coelomic cavity and accumulate in the coelom until spawning. The largest intraovarian oocytes were oval, up to 205x
<quantity id="4C889B19FFF4FFF9FB71F9D53B1CF9A3" box="[1205,1294,1581,1608]" metricMagnitude="-1" metricUnit="m" metricValue="2.75" pageId="14" pageNumber="267" unit="mm" value="275.0">275 µm</quantity>
in diameter, with the germinal vesicle about
<quantity id="4C889B19FFF4FFF9FDC7F9A93C5FF98F" box="[515,589,1617,1643]" metricMagnitude="-2" metricUnit="m" metricValue="8.0" pageId="14" pageNumber="267" unit="mm" value="80.0">80 µm</quantity>
and a single nucleolus
<quantity id="4C889B19FFF4FFF9FC9CF9A93DB0F98F" box="[856,930,1617,1644]" metricMagnitude="-2" metricUnit="m" metricValue="2.5" pageId="14" pageNumber="267" unit="mm" value="25.0">25 µm</quantity>
in diameter. The oocyte envelope was about
<quantity id="4C889B19FFF4FFF9FF53F98D3EF0F96B" box="[151,226,1653,1680]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="14" pageNumber="267" unit="mm" value="15.0">15 µm</quantity>
thick, with a honey-combed sculptured external surface, and pear-shaped cortical alveoli arranged in two parallel complete circles and one oblique incomplete row situated between the circles (
<figureCitation id="134B2A79FFF4FFF9FB9BF9613ACEF957" box="[1119,1244,1689,1715]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9AD</figureCitation>
). The large circle comprised 1417 alveoli situated close to the attachment place of the oocyte to the blood vessel (
<figureCitation id="134B2A79FFF4FFF9FB26F9453B4EF933" box="[1250,1372,1725,1751]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9B, C</figureCitation>
). The small circle comprised 79 alveoli (
<figureCitation id="134B2A79FFF4FFF9FDE6F9193C8BF91F" box="[546,665,1761,1787]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9A, B</figureCitation>
), while the intermediate row comprised about 10 alveoli (
<figureCitation id="134B2A79FFF4FFF9FAD2F9193B82F91F" box="[1302,1424,1761,1787]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9AC</figureCitation>
). Each alveolus was up to
<quantity id="4C889B19FFF4FFF9FE6DF8FD3FE3F8FB" box="[425,497,1797,1823]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="14" pageNumber="267" unit="mm" value="20.0">20 µm</quantity>
deep, with an external opening about
<quantity id="4C889B19FFF4FFF9FC5CF8FD3DF2F8FB" box="[920,992,1797,1823]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="14" pageNumber="267" unit="mm" value="10.0">10 µm</quantity>
in diameter, and an inner widest part up to
<quantity id="4C889B19FFF4FFF9FF70F8D13EEFF8A7" box="[180,253,1833,1859]" metricMagnitude="-2" metricUnit="m" metricValue="1.4" pageId="14" pageNumber="267" unit="mm" value="14.0">14 µm</quantity>
in diameter. When artificially released into sea water, the oocytes became spherical (
<figureCitation id="134B2A79FFF4FFF9FB72F8D13B02F8A7" box="[1206,1296,1833,1859]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9D</figureCitation>
).
</paragraph>
<paragraph id="8BCF36FCFFF4FFF9FF03F8B53FC3F837" blockId="14.[151,1437,1293,2039]" pageId="14" pageNumber="267">
Spermatogonia proliferate in the testes; spermatogenesis occurs in the coelomic cavity. Spermatids were joined in tetrads. Spermatozoa were ect-aquasperm with a small rounded acrosome, a spherical nucleus about 3 μm in di- ameter, spherical mitochondria, probably four in number and each one less than
<quantity id="4C889B19FFF4FFF9FBDBF86D3A48F84B" box="[1055,1114,1941,1967]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="14" pageNumber="267" unit="mm" value="1.0">1 µm</quantity>
in diameter, and a flagellum about
<quantity id="4C889B19FFF4FFF9FF18F8413F37F837" box="[220,293,1977,2003]" metricMagnitude="-2" metricUnit="m" metricValue="8.7" pageId="14" pageNumber="267" unit="mm" value="87.0">87 µm</quantity>
long (
<figureCitation id="134B2A79FFF4FFF9FEA9F8413FD7F837" box="[365,453,1977,2003]" captionStart="FIGURE 9" captionStartId="16.[151,250,991,1015]" captionTargetBox="[161,1425,181,967]" captionTargetId="figure-19@16.[161,1425,181,967]" captionTargetPageId="16" captionText="FIGURE 9. Laonice irinae n. sp. gamete and chaetal morphology. AC, large intraovarian oocytes: A, top view, showing three groups of alveoli; B, lateral view, showing three groups of alveoli; C, bottom view, showing large complete circle and intermediate incomplete row of alveoli. D, developed oocyte soon after being artificially released into seawater. E, coelomic sperm: diads of spermatocytes, tetrads of spermatids, and solitary spermatozoa with small rounded acrosome, spherical nucleus and spherical mitochondria. F, neuropodial hooded hooks and sabre chaeta from middle chaetiger. Abbreviations: al—cortical alveoli in oocyte envelope; in—intermediate oblique incomplete row of about ten alveoli; la—large complete circle of 1417 alveoli; sm—small complete circle of eight alveoli; st—stalk connecting oocyte to wall of blood vessel. AC, E, F—MIMB 39044 (paratypes); D—MIMB 39043 (paratype). AC, F—formalin-fixed specimen; D, E—live material. Scale bars: AD = 50 µm; E, F = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.5069852" httpUri="https://zenodo.org/record/5069852/files/figure.png" pageId="14" pageNumber="267">Fig. 9E</figureCitation>
).
</paragraph>
<paragraph id="8BCF36FCFFF4FFF8FF03F8253AE8FEFA" blockId="14.[151,1437,1293,2039]" lastBlockId="15.[151,1437,151,466]" lastPageId="15" lastPageNumber="268" pageId="14" pageNumber="267">
<emphasis id="B904EAEEFFF4FFF9FF03F8253FEDF813" bold="true" box="[199,511,2013,2039]" pageId="14" pageNumber="267">Methylene green staining.</emphasis>
Intensely stained upper parts of frontal surfaces of notopodial postchaetal lamellae from chaetiger 2 to approximately chaetiger 9 (
<figureCitation id="134B2A79FFF5FFF8FD10FF6F3D3DFF56" box="[724,815,151,178]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="15" pageNumber="268">Fig. 6G</figureCitation>
). Usually no staining on prostomium, peristomium and ventral side of chaetigers; rarely, intensely stained mid-longitudinal line in anterior half of prostomium and a spot posterior to occipital antenna, and diffusely stained dorsal and ventral sides of anterior parts of prostomium and peristomium (except the most frontal edge) (
<figureCitation id="134B2A79FFF5FFF8FD4FFEFB3CEFFEF9" box="[651,765,259,285]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="15" pageNumber="268">Fig. 6F, G</figureCitation>
demonstrates a rare case of intense staining).
</paragraph>
</subSubSection>
<subSubSection id="C36A6577FFF5FFF8FF03FEDF3FB1F8BA" pageId="15" pageNumber="268" type="discussion">
<paragraph id="8BCF36FCFFF5FFF8FF03FEDF3B08FE36" blockId="15.[151,1437,151,466]" pageId="15" pageNumber="268">
<emphasis id="B904EAEEFFF5FFF8FF03FEDF3F29FEA5" bold="true" box="[199,315,295,321]" pageId="15" pageNumber="268">Remarks.</emphasis>
The specimens herein referred to
<taxonomicName id="4C704D7FFFF5FFF8FD01FEDF3D3BFEA4" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[709,809,295,320]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF5FFF8FD01FEDF3D3BFEA4" box="[709,809,295,320]" italics="true" pageId="15" pageNumber="268">L. irinae</emphasis>
</taxonomicName>
<emphasis id="B904EAEEFFF5FFF8FCF7FEDF3D67FEA5" bold="true" box="[819,885,295,321]" pageId="15" pageNumber="268">
<taxonomicNameLabel id="A2375795FFF5FFF8FCF7FEDF3D67FEA5" box="[819,885,295,321]" pageId="15" pageNumber="268" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
had earlier been misidentified as
<taxonomicName id="4C704D7FFFF5FFF8FB38FEDF3B8EFEA5" authorityName=": Sikorski" authorityYear="2003" box="[1276,1436,295,321]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="bahusiensis">
<emphasis id="B904EAEEFFF5FFF8FB38FEDF3B8EFEA5" box="[1276,1436,295,321]" italics="true" pageId="15" pageNumber="268">L. bahusiensis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFE14B0DFFF5FFF8FF5BFEB33F2FFE81" author="Sikorski, A. V." box="[159,317,331,357]" pageId="15" pageNumber="268" pagination="406 - 419" refId="ref18229" refString="Sikorski, A. V. (2002) On distinguishing the morphologically close species, Laonice cirrata and L. bahusiensis (Polychaeta, Spionidae). Zoologicheskii Zhurnal, 81 (4), 406 - 419. [in Russian with English Summary]" type="journal article" year="2002">Sikorski 2002</bibRefCitation>
,
<bibRefCitation id="EFE14B0DFFF5FFF8FE83FEB33F91FE81" author="Sikorski, A. V." box="[327,387,331,357]" pageId="15" pageNumber="268" pagination="316 - 345" refId="ref18276" refString="Sikorski, A. V. (2003) Laonice (Polychaeta, Spionidae) in the Arctic and the North Atlantic. Sarsia, 88 (5), 316 - 345. https: // doi. org / 10.1080 / 00364820310002551" type="journal article" year="2003">2003</bibRefCitation>
). However, in
<taxonomicName id="4C704D7FFFF5FFF8FDE7FEB33CD2FE81" authorityName=": Sikorski" authorityYear="2003" box="[547,704,331,357]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="bahusiensis">
<emphasis id="B904EAEEFFF5FFF8FDE7FEB33CD2FE81" box="[547,704,331,357]" italics="true" pageId="15" pageNumber="268">L. bahusiensis</emphasis>
</taxonomicName>
the nuchal organs terminate 010 chaetigers before the last branchiate chaetiger (
<figureCitation id="134B2A79FFF5FFF8FE9EFE973FD1FE6D" box="[346,451,367,393]" captionStart="FIGURE 19" captionStartId="26.[151,250,1807,1831]" captionTargetBox="[151,1436,1323,1784]" captionTargetId="figure-84@26.[151,1436,1323,1784]" captionTargetPageId="26" captionText="FIGURE 19. Relationships between the arithmetic difference between number of last branchiate chaetiger and length of nuchal organs (in chaetiger number) (Br-NO) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis (empty circles) and Laonice irinae n. sp. (filled circles). B, Laonice grimaldii n. sp. (empty circles) and Laonice mediterranea n. sp. (filled circles)." figureDoi="http://doi.org/10.5281/zenodo.5069872" httpUri="https://zenodo.org/record/5069872/files/figure.png" pageId="15" pageNumber="268">Fig. 19A</figureCitation>
), and the dorsal transverse crests first appear on a chaetiger after the end of the nuchal organs, whereas in
<taxonomicName id="4C704D7FFFF5FFF8FEA9FE6B3FDFFE48" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[365,461,403,428]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF5FFF8FEA9FE6B3FDFFE48" box="[365,461,403,428]" italics="true" pageId="15" pageNumber="268">L. irinae</emphasis>
</taxonomicName>
<emphasis id="B904EAEEFFF5FFF8FE17FE6B3C03FE49" bold="true" box="[467,529,403,429]" pageId="15" pageNumber="268">
<taxonomicNameLabel id="A2375795FFF5FFF8FE17FE6B3C03FE49" box="[467,529,403,429]" pageId="15" pageNumber="268" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
the nuchal organs usually extend beyond the last branchiate chaetiger (
<figureCitation id="134B2A79FFF5FFF8FAECFE6B3B9DFE49" box="[1320,1423,403,429]" captionStart="FIGURE 19" captionStartId="26.[151,250,1807,1831]" captionTargetBox="[151,1436,1323,1784]" captionTargetId="figure-84@26.[151,1436,1323,1784]" captionTargetPageId="26" captionText="FIGURE 19. Relationships between the arithmetic difference between number of last branchiate chaetiger and length of nuchal organs (in chaetiger number) (Br-NO) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). A, Laonice bahusiensis (empty circles) and Laonice irinae n. sp. (filled circles). B, Laonice grimaldii n. sp. (empty circles) and Laonice mediterranea n. sp. (filled circles)." figureDoi="http://doi.org/10.5281/zenodo.5069872" httpUri="https://zenodo.org/record/5069872/files/figure.png" pageId="15" pageNumber="268">Fig. 19A</figureCitation>
), and the dorsal transverse crests first appear 15 chaetigers before the end of the nuchal organs (
<figureCitation id="134B2A79FFF5FFF8FB70FE4F3B1FFE36" box="[1204,1293,439,466]" captionStart="FIGURE 6" captionStartId="13.[151,250,1849,1873]" captionTargetBox="[154,1436,381,1826]" captionTargetId="figure-101@13.[151,1436,381,1826]" captionTargetPageId="13" captionText="FIGURE 6. Laonice irinae n. sp. adult morphology. A, anterior end, dorsal view. B, middle chaetigers, dorsal view, showing posterior end of caruncle with nuchal organs, and dorsal crests. CE, parapodia and branchiae, left sides, frontal view: C, chaetiger 4; D, chaetiger 5; E, chaetiger 6. F, G, anterior end, methylene green staining: ventral (F) and dorsal (G) view.A—ZMBN 135376; B—ZMBN 135384; CG—ZMBN 135383. Scale bars: A, B, F, G = 300 µm; CE = 200 µm." figureDoi="http://doi.org/10.5281/zenodo.5069846" httpUri="https://zenodo.org/record/5069846/files/figure.png" pageId="15" pageNumber="268">Fig. 6B</figureCitation>
).
</paragraph>
<caption id="DF0F6674FFF5FFF8FF53FA413B84F94D" ID-DOI="http://doi.org/10.5281/zenodo.5069850" ID-Zenodo-Dep="5069850" httpUri="https://zenodo.org/record/5069850/files/figure.png" pageId="15" pageNumber="268" startId="15.[151,250,1465,1489]" targetBox="[177,1411,489,1442]" targetPageId="15">
<paragraph id="8BCF36FCFFF5FFF8FF53FA413B84F94D" blockId="15.[151,1436,1465,1706]" pageId="15" pageNumber="268">
<emphasis id="B904EAEEFFF5FFF8FF53FA413FE9FA36" bold="true" box="[151,507,1465,1490]" pageId="15" pageNumber="268">
FIGURE 8.
<taxonomicName id="4C704D7FFFF5FFF8FED9FA413FAAFA35" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[285,440,1465,1489]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF5FFF8FED9FA413FAAFA35" bold="true" box="[285,440,1465,1489]" italics="true" pageId="15" pageNumber="268">Laonice irinae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2375795FFF5FFF8FE04FA423FE9FA36" box="[448,507,1466,1490]" pageId="15" pageNumber="268" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
adult morphometric relationships. A, length of nuchal organs (NO, in chaetiger numbers) versus width of chaetiger 7 in worm (filled circles), and distribution of branchiae (referring to the number of the last branchiate chaetiger) versus width of chaetiger 7 (empty circles). B, anterior position of sabre chaetae (referring to number of the first sabre-bearing chaetiger) versus width of chaetiger 7 (filled circles), and anterior position of hooks (referring to the number of the first hook-bearing chaetiger) versus width of chaetiger 7 (empty circles). C, length of nuchal organs (in chaetiger numbers) versus distribution of branchiae (referring to the number of the last branchiate chaetiger). D, anterior position of sabre chaetae versus distribution of branchiae (filled circles), and anterior position of hooks versus distribution of branchiae (empty circles).
</paragraph>
</caption>
<paragraph id="8BCF36FCFFF5FFF8FF03F9203FB1F8BA" blockId="15.[151,1437,1752,2030]" pageId="15" pageNumber="268">
Some individuals morphologically similar to
<taxonomicName id="4C704D7FFFF5FFF8FD00F9203D35F915" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[708,807,1752,1777]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF5FFF8FD00F9203D35F915" box="[708,807,1752,1777]" italics="true" pageId="15" pageNumber="268">L. irinae</emphasis>
</taxonomicName>
<emphasis id="B904EAEEFFF5FFF8FCF4F9203D63F916" bold="true" box="[816,881,1752,1778]" pageId="15" pageNumber="268">
<taxonomicNameLabel id="A2375795FFF5FFF8FCF4F9203D63F916" box="[816,881,1752,1778]" pageId="15" pageNumber="268" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
were collected on the shelf of
<collectingCountry id="F367766CFFF5FFF8FB17F9203B22F916" box="[1235,1328,1752,1778]" name="Portugal" pageId="15" pageNumber="268">Portugal</collectingCountry>
(material deposited at the Biological department of the University of
<collectingRegion id="49B4F81EFFF5FFF8FCE0F9043D63F8F2" box="[804,881,1788,1814]" country="Portugal" name="Aveiro" pageId="15" pageNumber="268">Aveiro</collectingRegion>
,
<collectingCountry id="F367766CFFF5FFF8FCB8F9043DCCF8F3" box="[892,990,1788,1815]" name="Portugal" pageId="15" pageNumber="268">Portugal</collectingCountry>
). Considering the long distance from the southern distributional limits of
<taxonomicName id="4C704D7FFFF5FFF8FE32F8D83C44F8DD" authority="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck, 2021" authorityName="Sikorski &amp; Radashevsky &amp; Castelli &amp; Pavlova &amp; Nygren &amp; Malyar &amp; Borisova &amp; Mikac &amp; Rousou &amp; Martin &amp; Gil &amp; Pacciardi &amp; Langeneck" authorityYear="2021" box="[502,598,1824,1849]" class="Polychaeta" family="Spionidae" genus="Laonice" kingdom="Animalia" order="Spionida" pageId="15" pageNumber="268" phylum="Annelida" rank="species" species="irinae" status="sp. nov.">
<emphasis id="B904EAEEFFF5FFF8FE32F8D83C44F8DD" box="[502,598,1824,1849]" italics="true" pageId="15" pageNumber="268">L. irinae</emphasis>
</taxonomicName>
<emphasis id="B904EAEEFFF5FFF8FD99F8D83C89F8DE" bold="true" box="[605,667,1824,1850]" pageId="15" pageNumber="268">
<taxonomicNameLabel id="A2375795FFF5FFF8FD99F8D83C89F8DE" box="[605,667,1824,1850]" pageId="15" pageNumber="268" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
reported here, and lacking molecular support, we decided to postpone their final identification.
</paragraph>
</subSubSection>
<subSubSection id="C36A6577FFF5FFF8FF03F8903A80F866" box="[199,1170,1896,1922]" pageId="15" pageNumber="268" type="etymology">
<paragraph id="8BCF36FCFFF5FFF8FF03F8903A80F866" blockId="15.[151,1437,1752,2030]" box="[199,1170,1896,1922]" pageId="15" pageNumber="268">
<emphasis id="B904EAEEFFF5FFF8FF03F8903F5CF866" bold="true" box="[199,334,1896,1922]" pageId="15" pageNumber="268">Etymology.</emphasis>
The species is named in honour of
<taxonomicName id="4C704D7FFFF5FFF8FD17F8903D99F866" authority="Sikorskaya" authorityName="Sikorskaya" box="[723,907,1896,1922]" class="Magnoliopsida" family="Sapindaceae" genus="Irina" higherTaxonomySource="CoL" kingdom="Plantae" order="Sapindales" pageId="15" pageNumber="268" phylum="Tracheophyta" rank="genus">Irina Sikorskaya</taxonomicName>
, wife of the first author.
</paragraph>
</subSubSection>
<subSubSection id="C36A6577FFF5FFF8FF03F8733FCFF80A" pageId="15" pageNumber="268" type="distribution">
<paragraph id="8BCF36FCFFF5FFF8FF03F8733FCFF80A" blockId="15.[151,1437,1752,2030]" pageId="15" pageNumber="268">
<emphasis id="B904EAEEFFF5FFF8FF03F8733F73F841" bold="true" box="[199,353,1931,1957]" pageId="15" pageNumber="268">Distribution.</emphasis>
<collectingCountry id="F367766CFFF5FFF8FEACF8743C11F842" box="[360,515,1932,1958]" name="Faroe Islands" pageId="15" pageNumber="268">Faroe Islands</collectingCountry>
; Shetland Islands; Norwegian Sea, from Bogelva (
<geoCoordinate id="EE44503BFFF5FFF8FBF3F8743A98F843" box="[1079,1162,1932,1959]" degrees="68.7" direction="north" orientation="latitude" pageId="15" pageNumber="268" precision="5555" value="68.7">68.7°N</geoCoordinate>
) south to the North Sea, Gullmarfjord,
<collectingCountry id="F367766CFFF5FFF8FEFCF8483F80F82E" box="[312,402,1968,1994]" name="Sweden" pageId="15" pageNumber="268">Sweden</collectingCountry>
, and Limfjord,
<collectingCountry id="F367766CFFF5FFF8FDF9F8483CBBF82E" box="[573,681,1968,1994]" name="Denmark" pageId="15" pageNumber="268">Denmark</collectingCountry>
; English Channel; Irish Sea, Liverpool Bay;
<collectingCountry id="F367766CFFF5FFF8FB5DF8483AF9F82E" box="[1177,1259,1968,1994]" name="Ireland" pageId="15" pageNumber="268">Ireland</collectingCountry>
, Deenish Island (
<figureCitation id="134B2A79FFF5FFF8FF64F82C3EEAF80A" box="[160,248,2003,2030]" captionStart="FIGURE 5" captionStartId="12.[151,250,1000,1024]" captionTargetBox="[151,1436,417,977]" captionTargetId="figure-107@12.[151,1436,417,977]" captionTargetPageId="12" captionText="FIGURE 5. Maps showing records of Laonice bahusiensis complex species in the North East Atlantic Ocean (see Tables 1, S1S2 for details). A, Laonice bahusiensis: red star—type locality: Skagerrak, Gullmarfjord, Bohuslän, Sweden; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology. B, Laonice irinae n. sp.: red star—type locality: Norwegian Sea, Mikkelsøya, Norway; green triangles—specimens sequenced in the present study; yellow circles—specimens identified based only on morphology." figureDoi="http://doi.org/10.5281/zenodo.5069844" httpUri="https://zenodo.org/record/5069844/files/figure.png" pageId="15" pageNumber="268">Fig. 5B</figureCitation>
). At
<quantity id="4C889B19FFF5FFF8FEE9F82B3F82F80A" box="[301,400,2003,2030]" metricMagnitude="1" metricUnit="m" metricValue="8.0" metricValueMax="15.6" metricValueMin="0.4" pageId="15" pageNumber="268" unit="m" value="80.0" valueMax="156.0" valueMin="4.0">4156 m</quantity>
depth.
</paragraph>
</subSubSection>
</treatment>
</document>