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<document id="46120ED246D84EF24149E56E84749100" ID-CLB-Dataset="308008" ID-DOI="10.1093/zoolinnean/zlae160" ID-GBIF-Dataset="f442baa6-585d-4612-803f-32db5f2baa5d" ID-ISSN="0024-4082" ID-Zenodo-Dep="14825615" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="valdenar" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="valdenar" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="valdenar" checkinTime="1738157910715" checkinUser="plazi" docAuthor="Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke &amp; TØttrup, Frederik LeerhØi Anders P." docDate="2024" docId="039F879BFF94FF854AD8F981FE1DF8B4" docLanguage="en" docName="zlae160.pdf" docOrigin="Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4)" docSource="https://doi.org/10.1093/zoolinnean/zlae160" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.10:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="10" docTitle="Hypsibius scabropygus Cuenot 1929" docType="treatment" docVersion="3" lastPageNumber="7" masterDocId="FFA6FFE3FF90FF83494BFFC0FF91FF94" masterDocTitle="Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae)" masterLastPageNumber="18" masterPageNumber="1" pageNumber="5" updateTime="1738851711255" updateUser="ExternalLinkService">
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<mods:title id="1BD2D15CDF853E2F998C8B50C9352F5D">Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae)</mods:title>
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<mods:namePart id="2460EFA261D12738476ECD929BF08D77">Gąsiorek, Piotr</mods:namePart>
<mods:affiliation id="B2989FB04DC084A4C045C36040C883FB">Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark &amp; Department of Invertebrate Evolution, Faculty of Biology, Jagiellonian University, Kraków, Poland</mods:affiliation>
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<mods:namePart id="E3DA96CFECFF25DB080184F407D564BE">SØrensen, Martin V.</mods:namePart>
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<mods:namePart id="7DD5E0B46970F6A1AFDB9A444F69C6CA">Lillemark, Marie Rathcke</mods:namePart>
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<mods:date id="FFB3A1022D694714F8715636A79EB339">2024</mods:date>
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<treatment id="039F879BFF94FF854AD8F981FE1DF8B4" ID-DOI="http://doi.org/10.5281/zenodo.14825617" ID-Zenodo-Dep="14825617" LSID="urn:lsid:plazi:treatment:039F879BFF94FF854AD8F981FE1DF8B4" httpUri="http://treatment.plazi.org/id/039F879BFF94FF854AD8F981FE1DF8B4" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5" scope_class="Eutardigrada" scope_family="Hypsibiidae" scope_order="Parachela" scope_phylum="Tardigrada" scope_superFamily="Hypsibioidea">
<subSubSection id="C32C6506FF94FF874AD8F981FAF9F9CF" box="[915,1384,1600,1627]" pageId="4" pageNumber="5" type="nomenclature">
<paragraph id="8B89368DFF94FF874AD8F981FAF9F9CF" blockId="4.[824,1477,1600,1977]" box="[915,1384,1600,1627]" pageId="4" pageNumber="5">
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Species:
<taxonomicName id="4C364D0EFF94FF874AA5F980FAF9F9CF" ID-CoL="3P3LG" authority="Cuenot, 1929" authorityName="Cuenot" authorityYear="1929" box="[1006,1384,1600,1627]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="4" pageNumber="5" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF94FF874AA5F980FB43F9CF" bold="true" box="[1006,1234,1600,1627]" italics="true" pageId="4" pageNumber="5">Hypsibius scabropygus</emphasis>
<bibRefCitation id="EFA74B7CFF94FF874D9DF980FAF9F9CF" author="Cuenot L" box="[1238,1384,1600,1627]" pageId="4" pageNumber="5" pagination="121 - 5" refId="ref10280" refString="Cuenot L. Description d'un tardigrade nouveau de la faune francaise. Archives d'Anatomie Microscopique 1929; 25: 121 - 5." type="journal article" year="1929">Cuénot, 1929</bibRefCitation>
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<paragraph id="8B89368DFF94FF864A72F9A7FDCCFB5D" blockId="4.[824,1477,1600,1977]" lastBlockId="5.[113,763,1076,1977]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
Body cylindrical, with weakly delineated and blunt cephalic region (
<figureCitation id="130D2A08FF94FF874AC6F947FC49F90B" box="[909,984,1671,1695]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="4" pageNumber="5">Fig. 3A</figureCitation>
). Eyes present as a group of pigment granules, which dissolve in Hoyers medium. Entire dorsal cuticle wrinkled (
<figureCitation id="130D2A08FF94FF874A3FF905FC2EF949" box="[884,959,1733,1757]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="4" pageNumber="5">Figs 4A</figureCitation>
,
<figureCitation id="130D2A08FF94FF874A81F905FB90F949" box="[970,1025,1733,1757]" captionStart="Figure 5" captionStartId="8.[129,194,1372,1396]" captionTargetBox="[161,1441,144,1344]" captionTargetId="figure-324@8.[161,1441,144,1344]" captionTargetPageId="8" captionText="Figure 5. Variability of cuticular sculpturing of Hypsibius scabropygus (all but B—Denmark, SEM; B—Poland, PCM): A, B, habitus in dorsal view; C, D, cephalic region; EH, caudal body part. Symbols: black arrowheads—oval muscle attachment sites, black empty arrowheads— longitudinal muscle attachment sites, asterisks—rudimentary elliptical organs.Scale bars = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.14825631" httpUri="https://zenodo.org/record/14825631/files/figure.png" pageId="4" pageNumber="5">5A, B</figureCitation>
) and with marked muscle attachment points in the middle of the trunk; posteriorly, the wrinkling changes into distinct sculpturing in the form of large, plaque-like protuberances in the caudal body portion (
<figureCitation id="130D2A08FF94FF874D8BF8E3FABCF8AF" box="[1216,1325,1827,1851]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="4" pageNumber="5">Figs 4BD</figureCitation>
,
<figureCitation id="130D2A08FF94FF874C71F8E3FA20F8AF" box="[1338,1457,1827,1851]" captionStart="Figure 5" captionStartId="8.[129,194,1372,1396]" captionTargetBox="[161,1441,144,1344]" captionTargetId="figure-324@8.[161,1441,144,1344]" captionTargetPageId="8" captionText="Figure 5. Variability of cuticular sculpturing of Hypsibius scabropygus (all but B—Denmark, SEM; B—Poland, PCM): A, B, habitus in dorsal view; C, D, cephalic region; EH, caudal body part. Symbols: black arrowheads—oval muscle attachment sites, black empty arrowheads— longitudinal muscle attachment sites, asterisks—rudimentary elliptical organs.Scale bars = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.14825631" httpUri="https://zenodo.org/record/14825631/files/figure.png" pageId="4" pageNumber="5">5A, B, EH</figureCitation>
). The protuberances are opalescent under stereomicroscope and in PCM. Most often, the sculpturing is arranged in two belts: more anterior with smaller protuberances (rarely shaped in a rectangular, plate-like manner;
<figureCitation id="130D2A08FF94FF874D3FF861FB2FF82D" box="[1140,1214,1953,1977]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="4" pageNumber="5">Fig. 4C</figureCitation>
), and posteriormost with large, multangular protuberances (
<figureCitation id="130D2A08FF95FF864898FBF4FDAEFBD8" box="[467,575,1076,1100]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="5" pageNumber="6">Figs 4BD</figureCitation>
,
<figureCitation id="130D2A08FF95FF864B00FBF4FDF7FBD8" box="[587,614,1076,1100]" captionStart="Figure 5" captionStartId="8.[129,194,1372,1396]" captionTargetBox="[161,1441,144,1344]" captionTargetId="figure-324@8.[161,1441,144,1344]" captionTargetPageId="8" captionText="Figure 5. Variability of cuticular sculpturing of Hypsibius scabropygus (all but B—Denmark, SEM; B—Poland, PCM): A, B, habitus in dorsal view; C, D, cephalic region; EH, caudal body part. Symbols: black arrowheads—oval muscle attachment sites, black empty arrowheads— longitudinal muscle attachment sites, asterisks—rudimentary elliptical organs.Scale bars = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.14825631" httpUri="https://zenodo.org/record/14825631/files/figure.png" pageId="5" pageNumber="6">5B</figureCitation>
). In SEM, the dorsoposterior sculpturing can be characterized as locally thickened wrinkling (
<figureCitation id="130D2A08FF95FF864856FBB3FE16FB1E" box="[285,391,1138,1163]" captionStart="Figure 5" captionStartId="8.[129,194,1372,1396]" captionTargetBox="[161,1441,144,1344]" captionTargetId="figure-324@8.[161,1441,144,1344]" captionTargetPageId="8" captionText="Figure 5. Variability of cuticular sculpturing of Hypsibius scabropygus (all but B—Denmark, SEM; B—Poland, PCM): A, B, habitus in dorsal view; C, D, cephalic region; EH, caudal body part. Symbols: black arrowheads—oval muscle attachment sites, black empty arrowheads— longitudinal muscle attachment sites, asterisks—rudimentary elliptical organs.Scale bars = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.14825631" httpUri="https://zenodo.org/record/14825631/files/figure.png" pageId="5" pageNumber="6">Fig. 5EH</figureCitation>
). Moreover, flattened structures are identifiable in SEM (not visible in PCM) and interpreted as rudimentary elliptical sensory organs (
<figureCitation id="130D2A08FF95FF8648AEFB71FDDCFB5D" box="[485,589,1201,1225]" captionStart="Figure 5" captionStartId="8.[129,194,1372,1396]" captionTargetBox="[161,1441,144,1344]" captionTargetId="figure-324@8.[161,1441,144,1344]" captionTargetPageId="8" captionText="Figure 5. Variability of cuticular sculpturing of Hypsibius scabropygus (all but B—Denmark, SEM; B—Poland, PCM): A, B, habitus in dorsal view; C, D, cephalic region; EH, caudal body part. Symbols: black arrowheads—oval muscle attachment sites, black empty arrowheads— longitudinal muscle attachment sites, asterisks—rudimentary elliptical organs.Scale bars = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.14825631" httpUri="https://zenodo.org/record/14825631/files/figure.png" pageId="5" pageNumber="6">Fig. 5C, D</figureCitation>
).
</paragraph>
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<paragraph id="8B89368DFF95FF864939FCBCFEC3FB90" blockId="5.[113,1444,892,1028]" pageId="5" pageNumber="6">
<emphasis id="B942EA9FFF95FF864939FCBCFF5BFC00" bold="true" box="[114,202,892,916]" pageId="5" pageNumber="6">Figure 2.</emphasis>
Phylogenetic relationships within
<taxonomicName id="4C364D0EFF95FF864B45FCBCFD10FC00" authorityName="Pilato" authorityYear="1969" box="[526,641,892,916]" class="Eutardigrada" family="Hypsibiidae" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="family">Hypsibiidae</taxonomicName>
in the Bayesian and maximum likelihood analyses (largely congruent topologies, see Supporting Information, S3 for the ML tree revealing incongruences and detailed ingroup topology of
<taxonomicName id="4C364D0EFF95FF864D7EFC58FB58FC24" authorityName="Pilato" authorityYear="1969" box="[1077,1225,920,944]" class="Eutardigrada" family="Hypsibiidae" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="subFamily" subFamily="Itaquasconinae">Itaquasconinae</taxonomicName>
). The scale refers to the Bayesian tree and represents substitutions per position; posterior probability values are provided above the nodes, whereas bootstrap values can be found below the nodes; *, maximal support; #, no support.
<taxonomicName id="4C364D0EFF95FF864B95FC10FC34FC7C" baseAuthorityName="Richters" baseAuthorityYear="1900" box="[734,933,976,1000]" class="Eutardigrada" family="Calohypsibiidae" genus="Calohypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="species" species="ornatus">
<emphasis id="B942EA9FFF95FF864B95FC10FC34FC7C" box="[734,933,976,1000]" italics="true" pageId="5" pageNumber="6">Calohypsibius ornatus</emphasis>
</taxonomicName>
constitutes an outgroup. Newly added species and populations are in bold.
</paragraph>
</caption>
<paragraph id="8B89368DFF95FF8649C6FB11FD85F8A8" blockId="5.[113,763,1076,1977]" pageId="5" pageNumber="6">
A pair of cribriform muscle attachment sites (
<bibRefCitation id="EFA74B7CFF95FF864B06FB11FD7EFB7D" author="Kihm J-H &amp; Zawierucha K &amp; Rho HS" box="[589,751,1233,1257]" pageId="5" pageNumber="6" refId="ref12088" refString="Kihm J-H, Zawierucha K, Rho HS et al. Homology of the head sensory structures between Heterotardigrada and Eutardigrada supported in a new species of water bear (Ramazzottiidae: Ramazzottius). Zoological Letters 2023; 9: 22. https: // doi. org / 10.1186 / s 40851 - 023 - 00221 - w" type="journal volume" year="2023">
Kihm
<emphasis id="B942EA9FFF95FF864BC0FB11FD23FB7D" box="[651,690,1233,1257]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
. 2023
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) flanking the mouth opening present (
<figureCitation id="130D2A08FF95FF864B48FB30FDDCFA9C" box="[515,589,1264,1288]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Fig. 6F</figureCitation>
). Six peribuccal lobes present around the mouth ring (
<figureCitation id="130D2A08FF95FF8648B2FACFFDD1FAB3" box="[505,576,1295,1319]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Fig. 6E</figureCitation>
), lamellae or papulae absent. Bucco-pharyngeal apparatus of the
<taxonomicName id="4C364D0EFF95FF864B39FAEFFD6BFAD2" authorityName="Binda and Pilato" authorityYear="1986" box="[626,762,1327,1350]" class="Eutardigrada" family="Hypsibiidae" genus="Ramazzottius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF95FF864B39FAEFFD6BFAD2" box="[626,762,1327,1350]" italics="true" pageId="5" pageNumber="6">Ramazzottius</emphasis>
</taxonomicName>
<typeStatus id="548D882FFF95FF86493AFA8EFF0EFAF2" box="[113,159,1358,1382]" pageId="5" pageNumber="6">type</typeStatus>
(
<figureCitation id="130D2A08FF95FF8649FFFA8EFEB5FAF2" box="[180,292,1358,1382]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Figs 3BD</figureCitation>
,
<figureCitation id="130D2A08FF95FF864878FA8EFEE0FAF2" box="[307,369,1358,1382]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">6AD</figureCitation>
). OCA absent (
<figureCitation id="130D2A08FF95FF864B6FFA8EFDE1FAF2" box="[548,624,1358,1382]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Fig. 6E</figureCitation>
). Apophyses for the insertion of stylet muscles (AISMs) asymmetrical with respect to the frontal plane: dorsal apophysis is shorter, higher, and stumpy (
<figureCitation id="130D2A08FF95FF8649B7FA6CFED8FA50" box="[252,329,1452,1476]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Fig. 3D</figureCitation>
), with a prominent caudal apex (
<figureCitation id="130D2A08FF95FF864BEEFA6CFF0EFA77" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Figs 6A, B, D</figureCitation>
,
<figureCitation id="130D2A08FF95FF8649E7FA0CFF74FA77" box="[172,229,1484,1508]" captionStart="Figure 7" captionStartId="10.[128,193,833,857]" captionTargetBox="[225,1377,144,806]" captionTargetId="figure-5@10.[225,1377,144,806]" captionTargetPageId="10" captionText="Figure 7. Bucco-pharyngeal apparatus of Hypsibius scabropygus (Denmark, SEM): A, buccal crown in dorsal view; B, buccal crown in dorsoposterior view (black arrowhead—the porous area); CF, posterior portion of the buccal tube and pharyngeal structures. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825635" httpUri="https://zenodo.org/record/14825635/files/figure.png" pageId="5" pageNumber="6">7A, B</figureCitation>
). Ventral apophysis slenderer (
<figureCitation id="130D2A08FF95FF864B60FA0BFDE6FA77" box="[555,631,1483,1507]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Fig. 3D</figureCitation>
), with more developed caudal processes (
<figureCitation id="130D2A08FF95FF8648D0FA2BFE77F997" box="[411,486,1515,1539]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Fig. 6C</figureCitation>
). Lateral porous areas present in the buccal crown (
<figureCitation id="130D2A08FF95FF8648CEF9CAFE43F9B6" box="[389,466,1546,1570]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Figs 6E</figureCitation>
,
<figureCitation id="130D2A08FF95FF8648ABF9CAFE6DF9B5" box="[480,508,1546,1569]" captionStart="Figure 7" captionStartId="10.[128,193,833,857]" captionTargetBox="[225,1377,144,806]" captionTargetId="figure-5@10.[225,1377,144,806]" captionTargetPageId="10" captionText="Figure 7. Bucco-pharyngeal apparatus of Hypsibius scabropygus (Denmark, SEM): A, buccal crown in dorsal view; B, buccal crown in dorsoposterior view (black arrowhead—the porous area); CF, posterior portion of the buccal tube and pharyngeal structures. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825635" httpUri="https://zenodo.org/record/14825635/files/figure.png" pageId="5" pageNumber="6">7B</figureCitation>
). Furcae unmodified, of the
<taxonomicName id="4C364D0EFF95FF8649D2F9E9FF68F9D5" authorityName="Ehrenberg" authorityYear="1848" box="[153,249,1577,1601]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF95FF8649D2F9E9FF68F9D5" box="[153,249,1577,1601]" italics="true" pageId="5" pageNumber="6">Hypsibius</emphasis>
</taxonomicName>
<typeStatus id="548D882FFF95FF8649B4F9EAFEBFF9D6" box="[255,302,1578,1602]" pageId="5" pageNumber="6">type</typeStatus>
, with well-developed condyles (
<figureCitation id="130D2A08FF95FF864B31F9E9FD58F9D5" box="[634,713,1577,1601]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Figs 3D</figureCitation>
,
<figureCitation id="130D2A08FF95FF864B9EF9E9FF14F9F4" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">6G, H</figureCitation>
). Buccal tube narrow, with a clear dorsal longitudinal wall thickening (
<figureCitation id="130D2A08FF95FF8649B8F9A8FE15F9EB" box="[243,388,1640,1664]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">Fig. 6A, B, D</figureCitation>
), posteriorly bent and terminating with pharyngeal apophyses (
<figureCitation id="130D2A08FF95FF8648D6F947FE7CF90B" box="[413,493,1671,1695]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Figs 3D</figureCitation>
,
<figureCitation id="130D2A08FF95FF8648B1F947FDA9F90B" box="[506,568,1671,1695]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">6AD</figureCitation>
). Pharynx circular (
<figureCitation id="130D2A08FF95FF864937F967FF75F92A" box="[124,228,1703,1727]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Fig. 3C, D</figureCitation>
), with only two granular macroplacoids (
<figureCitation id="130D2A08FF95FF864BC2F967FD64F92B" box="[649,757,1703,1727]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Figs 3BD</figureCitation>
,
<figureCitation id="130D2A08FF95FF86493AF906FF3EF94A" box="[113,175,1734,1758]" captionStart="Figure 6" captionStartId="9.[113,179,1112,1136]" captionTargetBox="[114,1458,144,1085]" captionTargetId="figure-447@9.[114,1458,144,1085]" captionTargetPageId="9" captionText="Figure 6. Bucco-pharyngeal apparatus and cephalic/peribuccal structures of Hypsibius scabropygus (Denmark, SEM): A, B, apparatus in dorsal view; C, apparatus in ventral view; D, apparatus in dorsolateral view; E, peribuccal region (16, peribuccal lobes, black arrowhead— the porous area); F, cribriform muscle attachment site; G, H, furca. Arrows indicate dorsal longitudinal thickening of the buccal tube wall. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825633" httpUri="https://zenodo.org/record/14825633/files/figure.png" pageId="5" pageNumber="6">6AD</figureCitation>
,
<figureCitation id="130D2A08FF95FF8649F6F906FF66F94A" box="[189,247,1734,1758]" captionStart="Figure 7" captionStartId="10.[128,193,833,857]" captionTargetBox="[225,1377,144,806]" captionTargetId="figure-5@10.[225,1377,144,806]" captionTargetPageId="10" captionText="Figure 7. Bucco-pharyngeal apparatus of Hypsibius scabropygus (Denmark, SEM): A, buccal crown in dorsal view; B, buccal crown in dorsoposterior view (black arrowhead—the porous area); CF, posterior portion of the buccal tube and pharyngeal structures. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825635" httpUri="https://zenodo.org/record/14825635/files/figure.png" pageId="5" pageNumber="6">7CF</figureCitation>
). Macroplacoids are of similar size or the first is only slightly longer than the second. Both macroplacoids with a central or subcentral constriction (
<figureCitation id="130D2A08FF95FF86489FF8C5FDABF889" box="[468,570,1797,1821]" captionStart="Figure 7" captionStartId="10.[128,193,833,857]" captionTargetBox="[225,1377,144,806]" captionTargetId="figure-5@10.[225,1377,144,806]" captionTargetPageId="10" captionText="Figure 7. Bucco-pharyngeal apparatus of Hypsibius scabropygus (Denmark, SEM): A, buccal crown in dorsal view; B, buccal crown in dorsoposterior view (black arrowhead—the porous area); CF, posterior portion of the buccal tube and pharyngeal structures. Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825635" httpUri="https://zenodo.org/record/14825635/files/figure.png" pageId="5" pageNumber="6">Fig. 7CF</figureCitation>
), but only the first constriction visible in PCM (
<figureCitation id="130D2A08FF95FF8648D0F8E4FD92F8A8" box="[411,515,1828,1852]" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Fig. 3BD</figureCitation>
).
</paragraph>
<paragraph id="8B89368DFF95FF8649C6F883FAD8FA31" blockId="5.[113,763,1076,1977]" lastBlockId="5.[810,1460,1076,1601]" pageId="5" pageNumber="6">
Claws
<taxonomicName id="4C364D0EFF95FF86499AF884FEC8F8CF" ID-CoL="76WW" authorityName="Binda and Pilato" authorityYear="1986" box="[209,345,1860,1883]" class="Eutardigrada" family="Hypsibiidae" genus="Ramazzottius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF95FF86499AF884FEC8F8CF" box="[209,345,1860,1883]" italics="true" pageId="5" pageNumber="6">Ramazzottius</emphasis>
</taxonomicName>
-like (
<emphasis id="B942EA9FFF95FF8648DCF884FE5BF8CF" box="[407,458,1860,1883]" italics="true" pageId="5" pageNumber="6">sensu</emphasis>
<bibRefCitation id="EFA74B7CFF95FF86489AF884FDFFF8CF" author="Tumanov DV" box="[465,622,1859,1883]" pageId="5" pageNumber="6" pagination="1 - 37" refId="ref14148" refString="Tumanov DV. Integrative redescription of Hypsibius pallidoides Pilato et al., 2011 (Eutardigrada: Hypsibioidea) with the erection of a new genus and discussion on the phylogeny of Hypsibiidae. European Journal of Taxonomy 2020; 681: 1 - 37. https: // doi. org / 10.5852 / ejt. 2020.681" type="journal article" year="2020">Tumanov 2020</bibRefCitation>
), with elongated primary branches of external/posterior claws and thinned connective portions in the form of light-refracting units (
<figureCitation id="130D2A08FF95FF864B9FF842FF3CF82D" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8CL</figureCitation>
). SEM microphotographs revealed that these units are particularly slender and flexible (
<figureCitation id="130D2A08FF95FF864DD8FBF4FAA8FBDF" box="[1171,1337,1076,1100]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8E, G, JL</figureCitation>
). Posterior claws are higher than external claws IIII. Accessory points on all primary branches evident and clearly divergent (
<figureCitation id="130D2A08FF95FF864CC6FBB3FCF7FB3E" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8CL</figureCitation>
). Pseudolunulae at claw bases present (
<figureCitation id="130D2A08FF95FF864DB4FB52FAEEFB3D" box="[1279,1407,1170,1194]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8D, H, I</figureCitation>
) but often poorly visible. Typically, pseudolunulae are most prominent under claws IV. Pulvini absent. Almost none of the bar
<typeStatus id="548D882FFF95FF864A61FB30FCF0FA9C" box="[810,865,1264,1288]" pageId="5" pageNumber="6">types</typeStatus>
defined for hypsibiids (
<bibRefCitation id="EFA74B7CFF95FF864D15FB30FABFFA9C" author="Gasiorek P &amp; Blagden B &amp; Morek W" box="[1118,1326,1264,1288]" pageId="5" pageNumber="6" refId="ref11032" refString="Gasiorek P, Blagden B, Morek W et al. What is a ' strong' synapomorphy? Redescriptions of Murray's type species and descriptions of new taxa challenge the systematics of Hypsibiidae (Eutardigrada: Parachela). Zoological Journal of the Linnean Society 2024; 202: zlad 151. https: // doi. org / 10.1093 / zoolinnean / zlad 151" type="journal volume" year="2024">
Gąsiorek
<emphasis id="B942EA9FFF95FF864D8AFB31FB7DFA9C" box="[1217,1260,1264,1288]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
. 2024
</bibRefCitation>
) are present (as described by
<bibRefCitation id="EFA74B7CFF95FF864A97FACFFBF4FAB3" author="Cuenot L" box="[988,1125,1295,1319]" pageId="5" pageNumber="6" pagination="121 - 5" refId="ref10280" refString="Cuenot L. Description d'un tardigrade nouveau de la faune francaise. Archives d'Anatomie Microscopique 1929; 25: 121 - 5." type="journal article" year="1929">Cuénot 1929</bibRefCitation>
,
<bibRefCitation id="EFA74B7CFF95FF864D39FACFFB37FAB3" author="Cuenot L" box="[1138,1190,1295,1319]" pageId="5" pageNumber="6" pagination="1 - 96" refId="ref10303" refString="Cuenot L. Tardigrades. In: Lechevalier P (ed.), Faune de France. Paris, France. 1932; 24: 1 - 96." type="journal article" year="1932">1932</bibRefCitation>
), but in some specimens, transverse basal bars on the legs IIII (
<figureCitation id="130D2A08FF95FF864D88FAEFFA81FAD3" box="[1219,1296,1327,1351]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8D</figureCitation>
) are developed asymmetrically and irregularly. These structures must lie under the cuticle as they are unidentifiable in SEM, and probably serve as muscle attachment points for retractor leg muscles.
</paragraph>
<paragraph id="8B89368DFF95FF864A0EFA6CFAA3F9D5" blockId="5.[810,1460,1076,1601]" pageId="5" pageNumber="6">
Eggs laid in exuviae (up to
<specimenCount id="9D30FD04FF95FF864D14FA6CFB26FA50" box="[1119,1207,1452,1476]" count="5" pageId="5" pageNumber="6" type="egg">five eggs</specimenCount>
per exuvia were found), but not smooth. Chorion with densely granulated surface (
<figureCitation id="130D2A08FF95FF864CC6FA0BFCD4F997" captionStart="Figure 3" captionStartId="6.[129,194,1386,1410]" captionTargetBox="[161,1441,144,1358]" captionTargetId="figure-256@6.[161,1441,144,1358]" captionTargetPageId="6" captionText="Figure 3. General morphology of H. scabropygus (Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus in toto (ventral view); D, bucco-pharyngeal apparatus in toto (lateral view, arrows indicate AISM); E, egg.Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="5" pageNumber="6">Fig. 3E</figureCitation>
), but without any processes. The granulation is developed already within the ovary during chorion formation in the females body. No specimens with spermatozoa were found.
</paragraph>
<paragraph id="8B89368DFF95FF864A90F9A1FA93F9EF" blockId="5.[808,1462,1633,1978]" box="[987,1282,1633,1659]" pageId="5" pageNumber="6">
<emphasis id="B942EA9FFF95FF864A90F9A1FA93F9EF" box="[987,1282,1633,1659]" italics="true" pageId="5" pageNumber="6">Comparison with related species</emphasis>
</paragraph>
<paragraph id="8B89368DFF95FF854A61F947FDDBF9B1" blockId="5.[808,1462,1633,1978]" lastBlockId="6.[128,778,1486,1824]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
<taxonomicName id="4C364D0EFF95FF864A61F947FC16F90B" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" box="[810,903,1671,1695]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="genus" status="sp. nov.">
<emphasis id="B942EA9FFF95FF864A61F947FC16F90B" box="[810,903,1671,1695]" italics="true" pageId="5" pageNumber="6">Hypsibius</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27157E4FF95FF864AC6F948FC49F90B" box="[909,984,1672,1695]" pageId="5" pageNumber="6" rank="species">sp. nov.</taxonomicNameLabel>
, the new species represented only by one population from the vicinity of Kolding (Jutland), is similar to
<taxonomicName id="4C364D0EFF95FF864CD1F967FC0AF94A" authorityName="Cuenot" authorityYear="1929" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF95FF864CD1F967FC0AF94A" italics="true" pageId="5" pageNumber="6">H. scabropygus</emphasis>
</taxonomicName>
in most characters, but differently developed caudal sculpturing, which is less sclerotized, without protuberances, and resembles folding of the cerebral cortex (
<figureCitation id="130D2A08FF95FF864D8DF8C5FAB9F889" box="[1222,1320,1797,1821]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="5" pageNumber="6">Fig. 4F, G</figureCitation>
), allows for a quickdelimitationunderalightmicroscope.Itsclawlight-refracting units seem to be weakly developed (
<figureCitation id="130D2A08FF95FF864DC7F883FB42F8CF" box="[1164,1235,1859,1883]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8A</figureCitation>
) when compared with those of
<taxonomicName id="4C364D0EFF95FF864A30F8A3FB99F8EF" authorityName="Cuenot" authorityYear="1929" box="[891,1032,1891,1915]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF95FF864A30F8A3FB99F8EF" box="[891,1032,1891,1915]" italics="true" pageId="5" pageNumber="6">H. scabropygus</emphasis>
</taxonomicName>
(
<figureCitation id="130D2A08FF95FF864D5DF8A3FBE9F8EF" box="[1046,1144,1891,1915]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="5" pageNumber="6">Fig. 8CL</figureCitation>
). Due to the scarcity of material, we refrain from formal description of this species. Importantly,
<typeStatus id="548D882FFF95FF864A61F862FC18F82E" box="[810,905,1954,1978]" pageId="5" pageNumber="6" type="paratype">paratypes</typeStatus>
of
<taxonomicName id="4C364D0EFF95FF864AE1F862FBC5F82D" authorityName="Ramazzotti and Maucci" authorityYear="1983" box="[938,1108,1953,1977]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="5" pageNumber="6" phylum="Tardigrada" rank="species" species="camelopardalis">
<emphasis id="B942EA9FFF95FF864AE1F862FBC5F82D" box="[938,1108,1953,1977]" italics="true" pageId="5" pageNumber="6">H. camelopardalis</emphasis>
</taxonomicName>
also have faint light-refracting units (
<figureCitation id="130D2A08FF96FF8549C0FA0EFF5EFA72" box="[139,207,1486,1510]" captionStart="Figure 8" captionStartId="10.[129,195,1873,1897]" captionTargetBox="[129,1473,974,1845]" captionTargetId="figure-47@10.[129,1473,974,1845]" captionTargetPageId="10" captionText="Figure 8. Claws of Hypsibius species from the scabropygus group (all but A and B, H. scabropygus, A, H. sp. nov., B, H. camelopardalis (paratype, microphotograph taken in Modena; all but B and I—Denmark, I—Scotland): A, claws I (PCM); B, claws II (PCM); C, D, claws II (PCM); E, claws II (SEM); F, claws III (PCM); G, claws III (SEM); H, I, claws IV (PCM); JL, claws IV (SEM). Symbols: white incised arrowheads—light-refracting unit of the primary branch, white empty arrowheads—pseudolunula, white flat arrowheads—transverse bars. Scale bars = 5 µm." figureDoi="http://doi.org/10.5281/zenodo.14825637" httpUri="https://zenodo.org/record/14825637/files/figure.png" pageId="6" pageNumber="7">Fig. 8B</figureCitation>
). The population of
<taxonomicName id="4C364D0EFF96FF8548D0FA0FFDF0FA72" authorityName="Ramazzotti and Maucci" authorityYear="1983" box="[411,609,1486,1510]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" isUncertain="true" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="camelopardalis">
<emphasis id="B942EA9FFF96FF8548D0FA0FFE3FFA72" box="[411,430,1487,1510]" italics="true" pageId="6" pageNumber="7">H</emphasis>
. cf.
<emphasis id="B942EA9FFF96FF85489EFA0EFDF0FA72" box="[469,609,1486,1510]" italics="true" pageId="6" pageNumber="7">camelopardalis</emphasis>
</taxonomicName>
from
<collectingCountry id="F321761DFF96FF854BD0FA0EFD56FA72" box="[667,711,1486,1510]" name="Italy" pageId="6" pageNumber="7">Italy</collectingCountry>
shows that the sculpturing can be formed as typical multangular plaques (
<figureCitation id="130D2A08FF96FF8549C0F9CDFF5FF9B1" box="[139,206,1549,1573]" captionStart="Figure 4" captionStartId="7.[113,178,1088,1112]" captionTargetBox="[114,1458,144,1061]" captionTargetId="figure-442@7.[114,1458,144,1061]" captionTargetPageId="7" captionText="Figure 4. Variability of cuticular sculpturing of Hypsibius species from the scabropygus group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all H. scabropygus, Denmark); E, caudal body part (H. cf. camelopardalis, Italy); F, G, caudal body part (H. sp.nov., Denmark). Scale bars are in µm." figureDoi="http://doi.org/10.5281/zenodo.14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="6" pageNumber="7">Fig. 4E</figureCitation>
), weakly elevated above the epicuticle.
</paragraph>
<caption id="DF496605FF96FF8549CAFAAAFB4CFA0B" ID-DOI="http://doi.org/10.5281/zenodo.14825625" ID-Zenodo-Dep="14825625" httpUri="https://zenodo.org/record/14825625/files/figure.png" pageId="6" pageNumber="7" startId="6.[129,194,1386,1410]" targetBox="[161,1441,144,1358]" targetPageId="6" targetType="figure">
<paragraph id="8B89368DFF96FF8549CAFAAAFB4CFA0B" blockId="6.[129,1450,1386,1439]" pageId="6" pageNumber="7">
<emphasis id="B942EA9FFF96FF8549CAFAAAFF49FA16" bold="true" box="[129,216,1386,1410]" pageId="6" pageNumber="7">Figure 3.</emphasis>
General morphology of
<taxonomicName id="4C364D0EFF96FF85488BFAABFDD9FA16" authorityName="Cuenot" authorityYear="1929" box="[448,584,1386,1410]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF96FF85488BFAABFDD9FA16" box="[448,584,1386,1410]" italics="true" pageId="6" pageNumber="7">H. scabropygus</emphasis>
</taxonomicName>
(Denmark, PCM): A, habitus in dorsolateral view; B, C, bucco-pharyngeal apparatus
<emphasis id="B942EA9FFF96FF854C3BFAABFA3BFA16" box="[1392,1450,1387,1410]" italics="true" pageId="6" pageNumber="7">in toto</emphasis>
(ventral view); D, bucco-pharyngeal apparatus
<emphasis id="B942EA9FFF96FF854B70FA47FDE4FA0A" box="[571,629,1415,1438]" italics="true" pageId="6" pageNumber="7">in toto</emphasis>
(lateral view, arrows indicate AISM); E, egg. Scale bars are in µm.
</paragraph>
</caption>
<paragraph id="8B89368DFF96FF8549D7F9EDFE1DF8B4" blockId="6.[128,778,1486,1824]" pageId="6" pageNumber="7">
Among the better-documented literature records,
<taxonomicName id="4C364D0EFF96FF854B82F9EDFF67F9F0" authorityName="Cuenot" authorityYear="1929" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" isUncertain="true" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF96FF854B82F9EDFD4CF9D0" box="[713,733,1581,1604]" italics="true" pageId="6" pageNumber="7">H</emphasis>
. cf.
<emphasis id="B942EA9FFF96FF8549CAF98CFF67F9F0" box="[129,246,1612,1636]" italics="true" pageId="6" pageNumber="7">scabropygus</emphasis>
</taxonomicName>
from the Baltic States (
<bibRefCitation id="EFA74B7CFF96FF8548A5F98CFD48F9F0" author="Zawierucha K &amp; Dziamiecki J &amp; Jakubowska N" box="[494,729,1612,1636]" pageId="6" pageNumber="7" pagination="81 - 105" refId="ref14496" refString="Zawierucha K, Dziamiecki J, Jakubowska N et al. New tardigrade records for the Baltic states with a description of Minibiotus formosus sp. n. (Eutardigrada, Macrobiotidae). ZooKeys 2014; 408: 81 - 105. https: // doi. org / 10.3897 / zookeys. 408.6612" type="journal article" year="2014">
Zawierucha
<emphasis id="B942EA9FFF96FF854B25F98CFD09F9F0" box="[622,664,1612,1636]" italics="true" pageId="6" pageNumber="7">et al</emphasis>
. 2014
</bibRefCitation>
) exhibits posterior bars and certainly represents another new species. Also
<taxonomicName id="4C364D0EFF96FF8549A7F94BFDF1F936" authority="Binda and Pilato, 1985" authorityName="Binda and Pilato" authorityYear="1985" box="[236,608,1674,1698]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="ragonesei">
<emphasis id="B942EA9FFF96FF8549A7F94BFEF6F936" box="[236,359,1675,1698]" italics="true" pageId="6" pageNumber="7">H. ragonesei</emphasis>
<bibRefCitation id="EFA74B7CFF96FF854824F94AFDF1F936" author="Binda MG &amp; Pilato G" box="[367,608,1674,1698]" pageId="6" pageNumber="7" pagination="245 - 8" refId="ref10079" refString="Binda MG, Pilato G. Hypsibius ragonesei, nuova specie di eutardigrado di Sicilia. Animalia 1985; 12: 245 - 8." type="journal article" year="1985">Binda and Pilato, 1985</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C364D0EFF96FF854BDCF94BFF7CF956" authority="Abe, 2004" authorityName="Abe" authorityYear="2004" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="stiliferus">
<emphasis id="B942EA9FFF96FF854BDCF94BFC98F936" box="[663,777,1674,1698]" italics="true" pageId="6" pageNumber="7">H. stiliferus</emphasis>
<bibRefCitation id="EFA74B7CFF96FF8549CAF96AFF7CF956" author="Abe W" box="[129,237,1706,1730]" pageId="6" pageNumber="7" pagination="957 - 62" refId="ref9678" refString="Abe W. A new species of the genus Hypsibius (Tardigrada: Parachela: Hypsibiidae) from Sakhalin Island, Far East Russia. Zoological Science 2004; 21: 957 - 62. https: // doi. org / 10.2108 / zsj. 21.957" type="journal article" year="2004">Abe, 2004</bibRefCitation>
</taxonomicName>
show posterior bars; all other bar
<typeStatus id="548D882FFF96FF854B29F96AFD08F956" box="[610,665,1706,1730]" pageId="6" pageNumber="7">types</typeStatus>
are absent in the
<taxonomicName id="4C364D0EFF96FF85498EF90AFEC8F975" authorityName="Cuenot" authorityYear="1929" box="[197,345,1737,1761]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF96FF85498EF90AFEC8F975" box="[197,345,1737,1761]" italics="true" pageId="6" pageNumber="7">H. scabropygus</emphasis>
</taxonomicName>
complex of spp. Given the morphological oddities of
<taxonomicName id="4C364D0EFF96FF8549BBF929FE13F894" authorityName="Cuenot" authorityYear="1929" box="[240,386,1768,1792]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF96FF8549BBF929FE13F894" box="[240,386,1768,1792]" italics="true" pageId="6" pageNumber="7">H. scabropygus</emphasis>
</taxonomicName>
and several other spp. (
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Parachela
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.,
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<emphasis id="B942EA9FFF96FF854A72FA0FFC32FA72" box="[825,931,1487,1510]" italics="true" pageId="6" pageNumber="7">Diagnosis:</emphasis>
Hypsibiids with
<taxonomicName id="4C364D0EFF96FF854DCAFA0FFA9CFA72" authorityName="Binda and Pilato" authorityYear="1986" box="[1153,1293,1487,1510]" class="Eutardigrada" family="Hypsibiidae" genus="Ramazzottius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF96FF854DCAFA0FFA9CFA72" box="[1153,1293,1487,1510]" italics="true" pageId="6" pageNumber="7">Ramazzottius</emphasis>
</taxonomicName>
-like claws and rigid buccal tube bent posteriorly. Only posterior bars may be present. AISMs asymmetrical, dorsal AISM shorter and higher than ventral, with prominent apex and less evident caudal processes. Pharynx with two granular macroplacoids. Rudimentary elliptical sensory organs present on head. Cuticular sculpturing well developed as protuberances at least in the dorsoposterior body portion, or protuberances cover an entire dorsum. Eggs laid within the exuviae, with granulated chorion.
</paragraph>
</subSubSection>
<subSubSection id="C32C6506FF96FF854A72F8DAFAD0F8C6" pageId="6" pageNumber="7" type="etymology">
<paragraph id="8B89368DFF96FF854A72F8DAFAD0F8C6" blockId="6.[825,1474,1818,1874]" pageId="6" pageNumber="7">
<emphasis id="B942EA9FFF96FF854A72F8DAFC39F8A6" box="[825,936,1818,1842]" italics="true" pageId="6" pageNumber="7">Etymology:</emphasis>
From Greek
<emphasis id="B942EA9FFF96FF854D75F8DBFBFAF8A6" box="[1086,1131,1819,1842]" italics="true" pageId="6" pageNumber="7">para</emphasis>
- = near, close +
<taxonomicName id="4C364D0EFF96FF854C56F8DAFAEDF8A6" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" box="[1309,1404,1818,1842]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF96FF854C56F8DAFAEDF8A6" box="[1309,1404,1818,1842]" italics="true" pageId="6" pageNumber="7">Hypsibius</emphasis>
</taxonomicName>
, referring to the close relationship between these genera.
</paragraph>
</subSubSection>
<subSubSection id="C32C6506FF96FF844A72F8B3FEB1FA00" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7" type="description">
<paragraph id="8B89368DFF96FF854A72F8B3FA53F85D" blockId="6.[825,1474,1907,1994]" pageId="6" pageNumber="7">
<emphasis id="B942EA9FFF96FF854A72F8B3FC24F81E" box="[825,949,1907,1930]" italics="true" pageId="6" pageNumber="7">Composition:</emphasis>
<taxonomicName id="4C364D0EFF96FF854A82F8B3FAB9F81F" authority="(Bartos, 1960)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Bartos" baseAuthorityYear="1960" box="[969,1320,1907,1931]" class="Eutardigrada" family="Hypsibiidae" genus="Parahypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="biscuitiformis" status="comb. nov.">
<emphasis id="B942EA9FFF96FF854A82F8B3FBE7F81F" bold="true" box="[969,1142,1907,1931]" italics="true" pageId="6" pageNumber="7">P. biscuitiformis</emphasis>
(
<bibRefCitation id="EFA74B7CFF96FF854DDAF8B3FAB0F81F" author="Bartos E" box="[1169,1313,1907,1931]" pageId="6" pageNumber="7" pagination="1 - 5" refId="ref9777" refString="Bartos E. Erganzungen zu der Tardigradenfauna Bohmens. Acta Universitatis Carolinae 1960; 1960: 1 - 5." type="journal article" year="1960">Bartoš, 1960</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF96FF854C77F8B3FA2BF81F" bold="true" box="[1340,1466,1907,1931]" pageId="6" pageNumber="7">
<taxonomicNameLabel id="A27157E4FF96FF854C77F8B3FA2BF81F" box="[1340,1466,1907,1931]" pageId="6" pageNumber="7" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
;
<taxonomicName id="4C364D0EFF96FF854A72F853FBF8F83E" authority="(Bartos, 1935)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Bartos" baseAuthorityYear="1935" box="[825,1129,1938,1962]" class="Eutardigrada" family="Hypsibiidae" genus="Parahypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="calcaratus" status="comb. nov.">
<emphasis id="B942EA9FFF96FF854A72F853FC50F83E" bold="true" box="[825,961,1938,1962]" italics="true" pageId="6" pageNumber="7">P. calcaratus</emphasis>
(
<bibRefCitation id="EFA74B7CFF96FF854A9CF852FBF3F83E" author="Bartos E" box="[983,1122,1938,1962]" pageId="6" pageNumber="7" pagination="257 - 60" refId="ref9725" refString="Bartos E. Vier neue Hypsibius - Arten aus der Tschechoslovakei. Zoologischer Anzeiger 1935; 110: 257 - 60." type="journal article" year="1935">Bartoš, 1935</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF96FF854D32F852FB63F83E" bold="true" box="[1145,1266,1938,1962]" pageId="6" pageNumber="7">
<taxonomicNameLabel id="A27157E4FF96FF854D32F852FB63F83E" box="[1145,1266,1938,1962]" pageId="6" pageNumber="7" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
;
<taxonomicName id="4C364D0EFF96FF854C42F853FBE9F85E" authority="(Ramazzotti and Maucci, 1983)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Ramazzotti and Maucci" baseAuthorityYear="1983" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="camelopardalis" status="comb. nov.">
<emphasis id="B942EA9FFF96FF854C42F853FA53F83E" bold="true" box="[1289,1474,1938,1962]" italics="true" pageId="6" pageNumber="7">P. camelopardalis</emphasis>
(
<bibRefCitation id="EFA74B7CFF96FF854A0BF872FBE0F85E" author="Ramazzotti G &amp; Maucci W" box="[832,1137,1970,1994]" pageId="6" pageNumber="7" pagination="1 - 1011" refId="ref13506" refString="Ramazzotti G, Maucci W. Il Phylum Tardigrada. III edizione riveduta e aggiornata. Memorie dell'Istituto Italiano di Idrobiologia 1983; 41: 1 - 1011." type="journal article" year="1983">Ramazzotti and Maucci, 1983</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF96FF854DCAF872FB62F85E" bold="true" box="[1153,1267,1970,1994]" pageId="6" pageNumber="7">
<taxonomicNameLabel id="A27157E4FF96FF854DCAF872FB62F85E" box="[1153,1267,1970,1994]" pageId="6" pageNumber="7" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
;
<taxonomicName id="4C364D0EFF96FF854C48F872FA53F85D" authorityName="Beasley" authorityYear="1988" box="[1283,1474,1969,1993]" class="Eutardigrada" family="Hypsibiidae" genus="Parahypsibius" kingdom="Animalia" order="Parachela" pageId="6" pageNumber="7" phylum="Tardigrada" rank="species" species="macrocalcaratus">
<emphasis id="B942EA9FFF96FF854C48F872FA53F85D" bold="true" box="[1283,1474,1969,1993]" italics="true" pageId="6" pageNumber="7">P. macrocalcaratus</emphasis>
</taxonomicName>
</paragraph>
<caption id="DF496605FF97FF84493AFB80FDE4FB05" ID-DOI="http://doi.org/10.5281/zenodo.14825629" ID-Zenodo-Dep="14825629" httpUri="https://zenodo.org/record/14825629/files/figure.png" pageId="7" pageNumber="8" startId="7.[113,178,1088,1112]" targetBox="[114,1458,144,1061]" targetPageId="7" targetType="figure">
<paragraph id="8B89368DFF97FF84493AFB80FDE4FB05" blockId="7.[113,1448,1088,1169]" pageId="7" pageNumber="8">
<emphasis id="B942EA9FFF97FF84493AFB80FF58FBCC" bold="true" box="[113,201,1088,1112]" pageId="7" pageNumber="8">Figure 4.</emphasis>
Variability of cuticular sculpturing of
<taxonomicName id="4C364D0EFF97FF844B67FB81FD5CFBCD" box="[556,717,1089,1113]" pageId="7" pageNumber="8">
<emphasis id="B942EA9FFF97FF844B67FB81FD14FBCD" box="[556,645,1089,1113]" italics="true" pageId="7" pageNumber="8">Hypsibius</emphasis>
species
</taxonomicName>
from the
<taxonomicName id="4C364D0EFF97FF844A62FB81FC04FBCD" authorityName="Cuenot" authorityYear="1929" box="[809,917,1089,1113]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF97FF844A62FB81FC04FBCD" box="[809,917,1089,1113]" italics="true" pageId="7" pageNumber="8">scabropygus</emphasis>
</taxonomicName>
group (PCM): A, frontal and middle body part; B, C, middle and caudal body part; D, caudal body part (all
<taxonomicName id="4C364D0EFF97FF844B26FB9DFD64FBE1" authorityName="Cuenot" authorityYear="1929" box="[621,757,1117,1141]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF97FF844B26FB9DFD64FBE1" box="[621,757,1117,1141]" italics="true" pageId="7" pageNumber="8">H. scabropygus</emphasis>
</taxonomicName>
, Denmark); E, caudal body part (
<taxonomicName id="4C364D0EFF97FF844D78FB9DFB60FBE1" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Ramazzotti and Maucci" baseAuthorityYear="1983" box="[1075,1265,1117,1141]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" isUncertain="true" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="camelopardalis">
<emphasis id="B942EA9FFF97FF844D78FB9DFBD4FBE0" box="[1075,1093,1117,1140]" italics="true" pageId="7" pageNumber="8">H</emphasis>
. cf.
<emphasis id="B942EA9FFF97FF844D21FB9DFB60FBE1" box="[1130,1265,1117,1141]" italics="true" pageId="7" pageNumber="8">camelopardalis</emphasis>
</taxonomicName>
, Italy); F, G, caudal body part (
<emphasis id="B942EA9FFF97FF844997FBB9FF7FFB04" box="[220,238,1145,1168]" italics="true" pageId="7" pageNumber="8">H</emphasis>
. sp. nov., Denmark). Scale bars are in µm.
</paragraph>
</caption>
<paragraph id="8B89368DFF97FF84493AFB00FEB1FA00" blockId="7.[113,764,1216,1428]" pageId="7" pageNumber="8">
(
<bibRefCitation id="EFA74B7CFF97FF844933FB00FE93FB4D" author="Beasley CW" box="[120,258,1216,1241]" pageId="7" pageNumber="8" pagination="436 - 40" refId="ref9797" refString="Beasley CW. Altitudinal distribution of Tardigrada of New Mexico with the description of a new species. American Midland Naturalist 1988; 120: 436 - 40. https: // doi. org / 10.2307 / 2426016" type="journal article" year="1988">Beasley, 1988</bibRefCitation>
)
<emphasis id="B942EA9FFF97FF84485BFB01FE11FB4D" bold="true" box="[272,384,1217,1241]" pageId="7" pageNumber="8">comb. nov.</emphasis>
;
<taxonomicName id="4C364D0EFF97FF8448C6FB01FD6AFB4C" authority="(Binda and Pilato, 1985)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Binda and Pilato" baseAuthorityYear="1985" box="[397,763,1216,1240]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="ragonesei" status="comb. nov.">
<emphasis id="B942EA9FFF97FF8448C6FB01FD93FB4C" bold="true" box="[397,514,1216,1240]" italics="true" pageId="7" pageNumber="8">P. ragonesei</emphasis>
(
<bibRefCitation id="EFA74B7CFF97FF844B44FB00FD65FB4C" author="Binda MG &amp; Pilato G" box="[527,756,1216,1240]" pageId="7" pageNumber="8" pagination="245 - 8" refId="ref10079" refString="Binda MG, Pilato G. Hypsibius ragonesei, nuova specie di eutardigrado di Sicilia. Animalia 1985; 12: 245 - 8." type="journal article" year="1985">Binda and Pilato, 1985</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF97FF844938FB20FF75FB6C" bold="true" box="[115,228,1248,1272]" pageId="7" pageNumber="8">
<taxonomicNameLabel id="A27157E4FF97FF844938FB20FF75FB6C" box="[115,228,1248,1272]" pageId="7" pageNumber="8" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
;
<taxonomicName id="4C364D0EFF97FF8449B8FB20FEFBFB6C" box="[243,362,1248,1272]" class="Eutardigrada" family="Hypsibiidae" genus="Parahypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="roanensis">
<emphasis id="B942EA9FFF97FF8449B8FB20FEFBFB6C" bold="true" box="[243,362,1248,1272]" italics="true" pageId="7" pageNumber="8">P. roanensis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFA74B7CFF97FF844832FB20FD3AFB6C" author="Nelson DR &amp; McGlothlin KL" box="[377,683,1248,1272]" pageId="7" pageNumber="8" pagination="140 - 4" refId="ref12682" refString="Nelson DR, McGlothlin KL. A new species of Hypsibius (Phylum Tardigrada) from Roan Mountain, Tennessee, U. S. A. Transactions of the American Microscopical Society 1993; 112: 140 - 4. https: // doi. org / 10.2307 / 3226827" type="journal article" year="1993">
<collectingRegion id="49F2F86FFF97FF844832FB20FE53FB6C" box="[377,450,1248,1272]" country="New Zealand" name="Nelson" pageId="7" pageNumber="8">Nelson</collectingRegion>
and McGlothlin, 1993
</bibRefCitation>
)
<emphasis id="B942EA9FFF97FF844BF1FB20FF0DFA83" bold="true" pageId="7" pageNumber="8">comb. nov.</emphasis>
;
<taxonomicName id="4C364D0EFF97FF8449E3FB3FFE47FA83" authority="(Cuenot, 1929)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Cuenot" baseAuthorityYear="1929" box="[168,470,1279,1303]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="scabropygus" status="comb. nov.">
<emphasis id="B942EA9FFF97FF8449E3FB3FFEA8FA83" bold="true" box="[168,313,1279,1303]" italics="true" pageId="7" pageNumber="8">P. scabropygus</emphasis>
(
<bibRefCitation id="EFA74B7CFF97FF84480FFB3FFE5EFA83" author="Cuenot L" box="[324,463,1279,1303]" pageId="7" pageNumber="8" pagination="121 - 5" refId="ref10280" refString="Cuenot L. Description d'un tardigrade nouveau de la faune francaise. Archives d'Anatomie Microscopique 1929; 25: 121 - 5." type="journal article" year="1929">Cuénot, 1929</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF97FF844891FB3FFDD8FA83" bold="true" box="[474,585,1279,1303]" pageId="7" pageNumber="8">
<taxonomicNameLabel id="A27157E4FF97FF844891FB3FFDD8FA83" box="[474,585,1279,1303]" pageId="7" pageNumber="8" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
;
<taxonomicName id="4C364D0EFF97FF844B1EFB3FFF3CFAA3" authority="(Abe, 2004)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Abe" baseAuthorityYear="2004" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="stiliferus" status="comb. nov.">
<emphasis id="B942EA9FFF97FF844B1EFB3FFD2EFA83" bold="true" box="[597,703,1279,1303]" italics="true" pageId="7" pageNumber="8">P. stiliferus</emphasis>
(
<bibRefCitation id="EFA74B7CFF97FF844B81FB3FFF37FAA3" author="Abe W" pageId="7" pageNumber="8" pagination="957 - 62" refId="ref9678" refString="Abe W. A new species of the genus Hypsibius (Tardigrada: Parachela: Hypsibiidae) from Sakhalin Island, Far East Russia. Zoological Science 2004; 21: 957 - 62. https: // doi. org / 10.2108 / zsj. 21.957" type="journal article" year="2004">Abe, 2004</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF97FF8449FEFADFFEB6FAA3" bold="true" box="[181,295,1311,1335]" pageId="7" pageNumber="8">
<taxonomicNameLabel id="A27157E4FF97FF8449FEFADFFEB6FAA3" box="[181,295,1311,1335]" pageId="7" pageNumber="8" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
; and
<taxonomicName id="4C364D0EFF97FF844829FADFFDDEFAA3" authority="(Bartos, 1941)" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Bartos" baseAuthorityYear="1941" box="[354,591,1311,1335]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="runae" status="comb. nov.">
<emphasis id="B942EA9FFF97FF844829FADFFE27FAA2" bold="true" box="[354,438,1311,1334]" italics="true" pageId="7" pageNumber="8">P. runae</emphasis>
(
<bibRefCitation id="EFA74B7CFF97FF84488FFADFFDD6FAA3" author="Bartos E" box="[452,583,1311,1335]" pageId="7" pageNumber="8" pagination="435 - 72" refId="ref9747" refString="Bartos E. Studien uber die Tardigraden des Karpathengebietes. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie der Tiere 1941; 74: 435 - 72." type="journal article" year="1941">Bartoš, 1941</bibRefCitation>
)
</taxonomicName>
<emphasis id="B942EA9FFF97FF844B1DFADFFD59FAA3" bold="true" box="[598,712,1311,1335]" pageId="7" pageNumber="8">
<taxonomicNameLabel id="A27157E4FF97FF844B1DFADFFD59FAA3" box="[598,712,1311,1335]" pageId="7" pageNumber="8" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
(uncertain,
<typeStatus id="548D882FFF97FF84498AFAFEFF7CFAC2" box="[193,237,1342,1366]" pageId="7" pageNumber="8">type</typeStatus>
specimens are unusable due to the crystallization of mounting medium, thus the transfer is done based on simplistic original description).
</paragraph>
</subSubSection>
<subSubSection id="C32C6506FF97FF84493AFA69FE1AF9AA" pageId="7" pageNumber="8" type="distribution">
<paragraph id="8B89368DFF97FF84493AFA69FE1AF9AA" blockId="7.[113,763,1449,1599]" pageId="7" pageNumber="8">
<emphasis id="B942EA9FFF97FF84493AFA69FECCFA55" box="[113,349,1449,1473]" italics="true" pageId="7" pageNumber="8">Geographic distribution:</emphasis>
Essentially Holarctic. Among all the species included in the genus, only
<taxonomicName id="4C364D0EFF97FF8448E4FA09FDD0FA74" authorityName="Gąsiorek &amp; SØrensen &amp; Lillemark &amp; TØttrup" authorityYear="2024" baseAuthorityName="Cuenot" baseAuthorityYear="1929" box="[431,577,1480,1504]" class="Eutardigrada" family="Hypsibiidae" genus="Hypsibius" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="scabropygus">
<emphasis id="B942EA9FFF97FF8448E4FA09FDD0FA74" box="[431,577,1480,1504]" italics="true" pageId="7" pageNumber="8">H. scabropygus</emphasis>
</taxonomicName>
was recorded outside of the Holarctic region—from the Palaeotropics (
<bibRefCitation id="EFA74B7CFF97FF844BEBFA28FF34F98B" author="McInnes SJ" pageId="7" pageNumber="8" pagination="257 - 352" refId="ref12489" refString="McInnes SJ. Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History 1994; 28: 257 - 352. https: // doi. org / 10.1080 / 00222939400770131" type="journal article" year="1994">McInnes 1994</bibRefCitation>
). The conspecificity of European and African populations has yet to be demonstrated.
</paragraph>
</subSubSection>
<subSubSection id="C32C6506FF97FF84493AF993FE76F893" pageId="7" pageNumber="8" type="diagnosis">
<paragraph id="8B89368DFF97FF84493AF993FE76F893" blockId="7.[113,764,1619,1800]" pageId="7" pageNumber="8">
<emphasis id="B942EA9FFF97FF84493AF993FED7F9FF" box="[113,326,1619,1643]" italics="true" pageId="7" pageNumber="8">Differential diagnosis:</emphasis>
The new genus is compared with all two-macroplacoid hypsibiid genera having a rigid buccal tube, and with
<taxonomicName id="4C364D0EFF97FF8449E1F952FD83F93E" authority="Pilato and Binda, 1997" authorityName="Pilato and Binda" authorityYear="1997" box="[170,530,1682,1706]" class="Eutardigrada" family="Hypsibiidae" genus="Acutuncus" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="genus">
<emphasis id="B942EA9FFF97FF8449E1F952FE9EF93D" box="[170,271,1682,1705]" italics="true" pageId="7" pageNumber="8">Acutuncus</emphasis>
Pilato and Binda, 1997
</taxonomicName>
(in
<bibRefCitation id="EFA74B7CFF97FF844B0DF952FF20F95D" author="Pilato G &amp; Binda MG" pageId="7" pageNumber="8" pagination="159 - 62" refId="ref13066" refString="Pilato G, Binda MG. Acutuncus, a new genus of Hypsibiidae (Eutardigrada). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 1997 b; 12: 159 - 62." type="journal article" year="1997">Pilato and Binda 1997b</bibRefCitation>
) due to the similar mode of egg deposition in the
<taxonomicName id="4C364D0EFF97FF84493AF911FEB0F97C" box="[113,289,1745,1768]" class="Eutardigrada" family="Hypsibiidae" genus="Acutuncus" kingdom="Animalia" order="Parachela" pageId="7" pageNumber="8" phylum="Tardigrada" rank="species" species="mariae">
<emphasis id="B942EA9FFF97FF84493AF911FEB0F97C" box="[113,289,1745,1768]" italics="true" pageId="7" pageNumber="8">Acutuncus mariae</emphasis>
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Vecchi
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<taxonomicName id="4C704D71FFFC0C23FF1A81B8FC93FC39" authority="Bloch &amp; Schneider, 1801" authorityName="Bloch &amp; Schneider" authorityYear="1801" box="[151,794,945,971]" class="Insecta" family="Brentidae" genus="Centrophorus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="510" phylum="Arthropoda" rank="species" species="granulosus">
<taxonomicName id="4C704D71FFFC0C23FF1A81B8FC93FC39" ID-CoL="69HMP" authority="(Bloch &amp; Schneider, 1801)" baseAuthorityName="Bloch &amp; Schneider" baseAuthorityYear="1801" box="[151,794,945,971]" class="Elasmobranchii" family="Centrophoridae" genus="Centrophorus" kingdom="Animalia" order="Squaliformes" pageId="3" pageNumber="510" phylum="Chordata" rank="species" species="granulosus">
<emphasis id="B904EAE0FFFC0C23FF1A81B8FC93FC39" bold="true" box="[151,794,945,971]" pageId="3" pageNumber="510">
<emphasis id="B904EAE0FFFC0C23FF1A81B8FE44FC39" bold="true" box="[151,461,945,971]" italics="true" pageId="3" pageNumber="510">Centrophorus granulosus</emphasis>
(Bloch &amp; Schneider, 1801)
@ -66,7 +67,7 @@ Bloch &amp; Schneider, 1801
locality: unknown (White
<emphasis id="B904EAE0FFFC0C23FB9981F0FBCEFBFC" box="[1044,1095,1016,1038]" italics="true" pageId="3" pageNumber="510">et al.</emphasis>
, 2013).
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<emphasis id="B904EAE0FFFC0C23FF1A861FFE10FBDE" box="[151,409,1046,1068]" italics="true" pageId="3" pageNumber="510">Centrophorus granulosus</emphasis>
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