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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: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">
<emphasis id="B942EA9FFF94FF874AD8F981FAF9F9CF" bold="true" box="[915,1384,1600,1627]" pageId="4" pageNumber="5">
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>
</taxonomicName>
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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>
<caption id="DF496605FF95FF864939FCBCFEC3FB90" ID-DOI="http://doi.org/10.5281/zenodo.14825621" ID-Zenodo-Dep="14825621" httpUri="https://zenodo.org/record/14825621/files/figure.png" pageId="5" pageNumber="6" startId="5.[114,179,892,916]" targetBox="[114,1452,148,859]" targetPageId="5" targetType="figure">
<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
</bibRefCitation>
) 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. (
<tableCitation id="C6B40336FF96FF854B3BF928FD2BF894" box="[624,698,1768,1792]" captionStart="Table 2" captionStartId="11.[113,169,143,167]" captionTargetPageId="11" captionText="Table 2. Comparison of the Hypsibius scabropygus complex vs. the Hypsibius dujardini and convergens morphogroups (the proposed Hypsibius s.s.)." httpUri="http://table.plazi.org/id/DF496605FF9BFF88493AFF4FFF6DFF57" pageId="6" pageNumber="7" tableUuid="DF496605FF9BFF88493AFF4FFF6DFF57">Table 2</tableCitation>
), a new hypsibiin genus is erected.
</paragraph>
</subSubSection>
</treatment>
</document>