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<mods:title id="F28FCEA6D73A680A1130C93497A8B699">Atlas of the Echiniscidae (Heterotardigrada) of the World-part I: West Palaearctic Echiniscus species</mods:title>
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<mods:affiliation id="EA9ABD7F2AB79593EE2DBA703EE6342E">Department of Invertebrate Evolution, Faculty of Biology, Jagiellonian University, Gronostajowa 9, 30 - 387 Kraków, Poland. &amp; Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5 - 7, DK- 1350; Universitetsparken 15, DK- 2100, Copenhagen, Denmark.</mods:affiliation>
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<mods:namePart id="EE3D24EE16DA299E149E645012BE7301">Vončina, Katarzyna</mods:namePart>
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<mods:affiliation id="1D085E45A3C60370833F0539560EF90B">Department of Invertebrate Evolution, Faculty of Biology, Jagiellonian University, Gronostajowa 9, 30 - 387 Kraków, Poland. &amp; Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt am Main, Germany. katarzyna. voncina @ senckenberg. de https: // orcid. org / 0000 - 0002 - 3210 - 0407 Correspondence: piotr. lukas. gasiorek @ gmail. com https: // orcid. org / 0000 - 0002 - 2814 - 8117 &amp; Department of Invertebrate Evolution, Faculty of Biology, Jagiellonian University, Gronostajowa 9, 30 - 387 Kraków, Poland.</mods:affiliation>
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<treatment id="8119D633B966FFDD1CEDFACCB8FCFF7B" ID-DOI="http://doi.org/10.5281/zenodo.8349268" ID-GBIF-Taxon="213153052" ID-Zenodo-Dep="8349268" LSID="urn:lsid:plazi:treatment:8119D633B966FFDD1CEDFACCB8FCFF7B" httpUri="http://treatment.plazi.org/id/8119D633B966FFDD1CEDFACCB8FCFF7B" lastPageId="42" lastPageNumber="42" pageId="39" pageNumber="40">
<subSubSection id="41AA34AEB966FFD01CEDFACCBBB6FAD0" box="[151,711,1380,1407]" pageId="39" pageNumber="40" type="nomenclature">
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<heading id="5247D049B966FFD01CEDFACCBBB6FAD0" bold="true" box="[151,711,1380,1407]" fontSize="11" level="2" pageId="39" pageNumber="40" reason="3">
<emphasis id="3BC4BB37B966FFD01CEDFACCBBB6FAD0" bold="true" box="[151,711,1380,1407]" pageId="39" pageNumber="40">
<emphasis id="3BC4BB37B966FFD01CEDFACCB9CDFAD0" bold="true" box="[151,188,1380,1406]" italics="true" pageId="39" pageNumber="40">15.</emphasis>
<taxonomicName id="CEB01CA6B966FFD01CBDFACCBBB6FAD0" ID-CoL="38G2Y" ID-ENA="1353343" authority="Richters, 1902" authorityName="Richters" authorityYear="1902" box="[199,711,1380,1407]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B966FFD01CBDFACCBB7DFAD1" bold="true" box="[199,524,1380,1407]" italics="true" pageId="39" pageNumber="40">Echiniscus quadrispinosus</emphasis>
<bibRefCitation id="6D211AD4B966FFD01E6EFACCBBB6FAD0" author="Richters, F." box="[532,711,1380,1406]" pageId="39" pageNumber="40" pagination="23 - 26" refId="ref37172" refString="Richters, F. (1902) Neue Moosbewohner. Bericht ¸ ber die Senckenbergische Naturforschende Gesselschaft in Frankfurt am Main, 23 - 26." type="book chapter" year="1902">Richters, 1902</bibRefCitation>
</taxonomicName>
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<paragraph id="090F6725B966FFD01CEDFA20BBE3FA0D" blockId="39.[151,711,1380,1443]" box="[151,658,1416,1443]" pageId="39" pageNumber="40">
<figureCitation id="918B7BA0B966FFD01CEDFA20B84AFA0D" box="[151,315,1416,1443]" captionStart-0="FIGURE 22" captionStart-1="FIGURE 23" captionStart-2="FIGURE 24" captionStart-3="FIGURE 25" captionStartId-0="39.[152,255,986,1010]" captionStartId-1="40.[152,255,1943,1967]" captionStartId-2="41.[152,255,1310,1334]" captionStartId-3="42.[152,255,1345,1369]" captionTargetBox-0="[151,1435,166,970]" captionTargetBox-1="[332,1255,181,1919]" captionTargetBox-2="[151,1435,181,1286]" captionTargetBox-3="[151,1435,222,1330]" captionTargetId-0="figure-19@39.[151,1435,166,970]" captionTargetId-1="figure-26@40.[332,1255,181,1919]" captionTargetId-2="figure-19@41.[151,1435,181,1286]" captionTargetId-3="figure-46@42.[151,1435,222,1330]" captionTargetPageId-0="39" captionTargetPageId-1="40" captionTargetPageId-2="41" captionTargetPageId-3="42" captionText-0="FIGURE 22. Echiniscus quadrispinosus from Germany (females): A—dorsal view (PCM), B—dorsal view (SEM), C— dorsolateral view (PCM), D—lateral view (SEM). Scale bars = 50 μm." captionText-1="FIGURE 23. Echiniscus quadrispinosus from Germany (PCM): A—male in dorsolateral view, B—juvenile in dorsolateral view, C—larva in dorsal view. Scale bars = 50 μm." captionText-2="FIGURE 24. Echiniscus quadrispinosus from Germany—cephalic region:A—scapular plate sculpturing in upper magnification (PCM), B—scapular plate sculpturing in lower magnification (PCM), C—scapular plate sculpturing (SEM), D—cephalic appendages and subcephalic plates (SEM). Scale bars = 20 μm." captionText-3="FIGURE 25. Echiniscus quadrispinosus from Germany—morphological details: A—claws III (PCM), B —claws III (SEM), C—claws IV (SEM), D—genital plates and gonopore (SEM). Scale bars = 10 μm." figureDoi-0="http://doi.org/10.5281/zenodo.8346101" figureDoi-1="http://doi.org/10.5281/zenodo.8346103" figureDoi-2="http://doi.org/10.5281/zenodo.8346105" figureDoi-3="http://doi.org/10.5281/zenodo.8346107" httpUri-0="https://zenodo.org/record/8346101/files/figure.png" httpUri-1="https://zenodo.org/record/8346103/files/figure.png" httpUri-2="https://zenodo.org/record/8346105/files/figure.png" httpUri-3="https://zenodo.org/record/8346107/files/figure.png" pageId="39" pageNumber="40">Figures 22–25</figureCitation>
, Supplementary Materials 5–8
</paragraph>
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<paragraph id="090F6725B966FFD01CEDFA65B873FA4B" blockId="39.[151,981,1485,1662]" box="[151,258,1485,1509]" pageId="39" pageNumber="40">Synonyms</paragraph>
<paragraph id="090F6725B966FFD01CEDFA44BBF8F9AA" blockId="39.[151,981,1485,1662]" box="[151,649,1515,1540]" pageId="39" pageNumber="40">
<taxonomicName id="CEB01CA6B966FFD01CEDFA44B89DF9AD" authority="C. A. S. Schultze, 1861" authorityName="C. A. S. Schultze" authorityYear="1861" box="[151,492,1515,1540]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="species" species="creplini">
<emphasis id="3BC4BB37B966FFD01CEDFA44B870F9AD" box="[151,257,1515,1539]" italics="true" pageId="39" pageNumber="40">E. creplini</emphasis>
C.A.S.
<bibRefCitation id="6D211AD4B966FFD01D28FA43B89DF9AD" author="Schultze, C. A. S." box="[338,492,1515,1539]" pageId="39" pageNumber="40" refId="ref37654" refString="Schultze, C. A. S. (1861) Echiniscus Creplini, Animalculum e Familia Arctiscoidum. Gryphiae, F. G., Kunike, 12 pp." type="book" year="1861">Schultze, 1861</bibRefCitation>
</taxonomicName>
:
<bibRefCitation id="6D211AD4B966FFD01D8CFA44BBF8F9AA" author="Cuenot, L." box="[502,649,1516,1540]" pageId="39" pageNumber="40" pagination="1 - 96" refId="ref29878" refString="Cuenot, L. (1932) Tardigrades. In: Lechevalier, P. (Ed.), Faune de France, 24, pp. 1 - 96." type="journal article" year="1932">Cuénot (1932)</bibRefCitation>
</paragraph>
<paragraph id="090F6725B966FFD01CEDF9A2BAA4F98C" blockId="39.[151,981,1485,1662]" box="[151,981,1546,1570]" pageId="39" pageNumber="40">
<taxonomicName id="CEB01CA6B966FFD01CEDF9A2BBA4F98C" ID-CoL="5H27Q" authority="Bartos, 1934" authorityName="Bartos" authorityYear="1934" box="[151,725,1546,1570]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="subSpecies" species="quadrispinosus" subSpecies="brachyspinosus">
<emphasis id="3BC4BB37B966FFD01CEDF9A2BB3BF98C" box="[151,586,1546,1570]" italics="true" pageId="39" pageNumber="40">Echiniscus quadrispinosus brachyspinosus</emphasis>
<bibRefCitation id="6D211AD4B966FFD01E2AF9A2BBA4F98C" author="Bartos, E." box="[592,725,1546,1570]" pageId="39" pageNumber="40" pagination="99 - 107" refId="ref28588" refString="Bartos, E. (1934) Prispevek k poznani Tardigrad CSR. Casopis Ceskoslovenske Spolecnosti Entomologicke, 31, 99 - 107." type="journal article" year="1934">Bartoš, 1934</bibRefCitation>
</taxonomicName>
:
<bibRefCitation id="6D211AD4B966FFD01EA5F9A2BAA4F98C" author="Kaczmarek, L. &amp; Kayastha, P. &amp; Gawlak, M. &amp; Mioduchowska, M. &amp; Roszkowska, M." box="[735,981,1546,1570]" pageId="39" pageNumber="40" pagination="102 - 124" refId="ref33566" refString="Kaczmarek, L., Kayastha, P., Gawlak, M., Mioduchowska, M. &amp; Roszkowska, M. (2022) An integrative redescription of Echiniscus quadrispinosus quadrispinosus Richters, 1902 (Heterotardigrada, Echiniscidae) from the terra typica in Taunus Mountain Range (Europe; Germany). European Journal of Taxonomy, 823 (1), 102 - 124. https: // doi. org / 10.5852 / ejt. 2022.823.1819" type="journal article" year="2022">
Kaczmarek
<emphasis id="3BC4BB37B966FFD01F23F9A3BAF7F98C" box="[857,902,1546,1570]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
. (2022)
</bibRefCitation>
</paragraph>
<paragraph id="090F6725B966FFD01CEDF981BAEEF9EF" blockId="39.[151,981,1485,1662]" box="[151,927,1577,1601]" pageId="39" pageNumber="40">
<taxonomicName id="CEB01CA6B966FFD01CEDF981BBEEF9EF" ID-CoL="5H27R" authority="Murray, 1907" authorityName="Murray" authorityYear="1907" box="[151,671,1577,1601]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="subSpecies" species="quadrispinosus" subSpecies="cribrosus">
<emphasis id="3BC4BB37B966FFD01CEDF981BB7AF9EF" box="[151,523,1577,1601]" italics="true" pageId="39" pageNumber="40">Echiniscus quadrispinosus cribrosus</emphasis>
Murray, 1907
</taxonomicName>
:
<bibRefCitation id="6D211AD4B966FFD01ED3F981BAEEF9EF" author="Kaczmarek, L. &amp; Kayastha, P. &amp; Gawlak, M. &amp; Mioduchowska, M. &amp; Roszkowska, M." box="[681,927,1577,1601]" pageId="39" pageNumber="40" pagination="102 - 124" refId="ref33566" refString="Kaczmarek, L., Kayastha, P., Gawlak, M., Mioduchowska, M. &amp; Roszkowska, M. (2022) An integrative redescription of Echiniscus quadrispinosus quadrispinosus Richters, 1902 (Heterotardigrada, Echiniscidae) from the terra typica in Taunus Mountain Range (Europe; Germany). European Journal of Taxonomy, 823 (1), 102 - 124. https: // doi. org / 10.5852 / ejt. 2022.823.1819" type="journal article" year="2022">
Kaczmarek
<emphasis id="3BC4BB37B966FFD01F58F982BA3EF9EF" box="[802,847,1577,1601]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
. (2022)
</bibRefCitation>
</paragraph>
<paragraph id="090F6725B966FFD01CEDF9EFBAB1F9CE" blockId="39.[151,981,1485,1662]" box="[151,960,1607,1632]" pageId="39" pageNumber="40">
<taxonomicName id="CEB01CA6B966FFD01CEDF9EFBBB1F9CE" ID-CoL="5H27S" authority="Murray, 1907" authorityName="Murray" authorityYear="1907" box="[151,704,1607,1632]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="subSpecies" species="quadrispinosus" subSpecies="fissispinosus">
<emphasis id="3BC4BB37B966FFD01CEDF9EFBB5DF9CE" box="[151,556,1607,1632]" italics="true" pageId="39" pageNumber="40">Echiniscus quadrispinosus fissispinosus</emphasis>
Murray, 1907
</taxonomicName>
:
<bibRefCitation id="6D211AD4B966FFD01EB0F9EFBAB1F9CE" author="Kaczmarek, L. &amp; Kayastha, P. &amp; Gawlak, M. &amp; Mioduchowska, M. &amp; Roszkowska, M." box="[714,960,1607,1632]" pageId="39" pageNumber="40" pagination="102 - 124" refId="ref33566" refString="Kaczmarek, L., Kayastha, P., Gawlak, M., Mioduchowska, M. &amp; Roszkowska, M. (2022) An integrative redescription of Echiniscus quadrispinosus quadrispinosus Richters, 1902 (Heterotardigrada, Echiniscidae) from the terra typica in Taunus Mountain Range (Europe; Germany). European Journal of Taxonomy, 823 (1), 102 - 124. https: // doi. org / 10.5852 / ejt. 2022.823.1819" type="journal article" year="2022">
Kaczmarek
<emphasis id="3BC4BB37B966FFD01F3EF9E0BA01F9F1" box="[836,880,1607,1631]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
. (2022)
</bibRefCitation>
</paragraph>
<paragraph id="090F6725B966FFD01CEDF9CFBBB3F9D0" blockId="39.[151,981,1485,1662]" box="[151,706,1638,1662]" pageId="39" pageNumber="40">
<taxonomicName id="CEB01CA6B966FFD01CEDF9CFB8FEF9D0" authority="Richters, 1902" authorityName="Richters" authorityYear="1902" box="[151,399,1638,1662]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="species" species="scrofa">
<emphasis id="3BC4BB37B966FFD01CEDF9CFB980F9D0" box="[151,241,1638,1662]" italics="true" pageId="39" pageNumber="40">E. scrofa</emphasis>
<bibRefCitation id="6D211AD4B966FFD01C82F9CEB8FEF9D0" author="Richters, F." box="[248,399,1638,1662]" pageId="39" pageNumber="40" pagination="23 - 26" refId="ref37172" refString="Richters, F. (1902) Neue Moosbewohner. Bericht ¸ ber die Senckenbergische Naturforschende Gesselschaft in Frankfurt am Main, 23 - 26." type="book chapter" year="1902">Richters, 1902</bibRefCitation>
</taxonomicName>
:
<bibRefCitation id="6D211AD4B966FFD01DE3F9CEBBB3F9D0" author="Ramazzotti, G. &amp; Maucci, W." box="[409,706,1638,1662]" pageId="39" pageNumber="40" pagination="1 - 1011" refId="ref37084" refString="Ramazzotti, G. &amp; Maucci, W. (1983) Il Phylum Tardigrada. III edizione riveduta e aggiornata. Memorie dell'Istituto Italiano di Idrobiologia, 41, 1 - 1011." type="journal article" year="1983">Ramazzotti &amp; Maucci (1983)</bibRefCitation>
</paragraph>
</subSubSection>
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<taxonomicName id="CEB01CA6B966FFD01CEDF905B86EF968" box="[151,287,1709,1734]" class="Insecta" family="Lycaenidae" genus="Terra" kingdom="Animalia" order="Lepidoptera" pageId="39" pageNumber="40" phylum="Arthropoda" rank="species" species="typica">
<emphasis id="3BC4BB37B966FFD01CEDF905B86EF968" box="[151,287,1709,1734]" italics="true" pageId="39" pageNumber="40">Terra typica</emphasis>
</taxonomicName>
:
<collectingCountry id="71A727B5B966FFD01D55F904B8E9F968" box="[303,408,1708,1734]" name="Germany" pageId="39" pageNumber="40">Germany</collectingCountry>
, Taunus Mountains.
</paragraph>
<paragraph id="090F6725B966FFD01CBDF978B8E4F8A0" blockId="39.[151,1436,1708,1984]" pageId="39" pageNumber="40">
<materialsCitation id="B9D86D78B966FFD01CBDF978B8E0F8A0" ID-GBIF-Occurrence="4411079309" country="Germany" elevation="485" latitude="50.196945" location="Konigstein im Taunus" longLatPrecision="18" longitude="8.454166" municipality="Taunus Mountains" pageId="39" pageNumber="40" specimenCount="1" stateProvince="Hessen">
New
<typeStatus id="D60BD987B966FFD01D73F978B848F944" box="[265,313,1744,1770]" pageId="39" pageNumber="40">type</typeStatus>
locality:
<geoCoordinate id="6C8401E2B966FFD01DD4F967BB4DF944" box="[430,572,1743,1770]" degrees="50" direction="north" minutes="11" orientation="latitude" pageId="39" pageNumber="40" precision="15" seconds="49" value="50.196945">50°11’49’’N</geoCoordinate>
,
<geoCoordinate id="6C8401E2B966FFD01E34F967BBBDF944" box="[590,716,1743,1770]" degrees="8" direction="east" minutes="27" orientation="longitude" pageId="39" pageNumber="40" precision="15" seconds="15" value="8.454166">8°27’15’’E</geoCoordinate>
,
<elevation id="829D8016B966FFD01EA4F967BA2BF944" box="[734,858,1743,1770]" metricMagnitude="2" metricUnit="m" metricValue="4.85" pageId="39" pageNumber="40" unit="m" value="485.0">
<quantity id="CE48CAC0B966FFD01EA4F967BA5BF944" box="[734,810,1743,1770]" metricMagnitude="2" metricUnit="m" metricValue="4.85" pageId="39" pageNumber="40" unit="m" value="485.0">485 m</quantity>
asl
</elevation>
:
<collectingCountry id="71A727B5B966FFD01F10F978BAA2F944" box="[874,979,1744,1770]" name="Germany" pageId="39" pageNumber="40">Germany</collectingCountry>
,
<collectingRegion id="CB74A9C7B966FFD01F9EF978BD5BF944" box="[996,1066,1744,1770]" country="Germany" name="Hessen" pageId="39" pageNumber="40">Hesse</collectingRegion>
,
<collectingMunicipality id="E96BFD5FB966FFD01840F978BC63F944" box="[1082,1298,1744,1770]" pageId="39" pageNumber="40">Taunus Mountains</collectingMunicipality>
, vicinity of
<location id="0C6F31FEB966FFD01CEDF95BB8E0F8A0" LSID="urn:lsid:plazi:treatment:8119D633B966FFDD1CEDFACCB8FCFF7B:0C6F31FEB966FFD01CEDF95BB8E0F8A0" box="[151,401,1779,1806]" country="Germany" latitude="50.196945" longLatPrecision="18" longitude="8.454166" municipality="Taunus Mountains" name="Konigstein im Taunus" pageId="39" pageNumber="40" stateProvince="Hessen">Königstein im Taunus</location>
</materialsCitation>
.
</paragraph>
<paragraph id="090F6725B966FFD01CBDF8BFBBBBF8D7" blockId="39.[151,1436,1708,1984]" pageId="39" pageNumber="40">
Additional localities: See
<tableCitation id="4432529EB966FFD01D88F8BFBB39F89F" box="[498,584,1815,1841]" captionStart="TABLE 1" captionStartId="5.[151,243,471,497]" captionTargetBox="[165,2025,548,1678]" captionTargetPageId="5" captionText="TABLE 1. List of sequenced populations of various European echiniscids. Types of analyses: (PCM) imaging and morphometry in PCM, (SEM) imaging in SEM, (DNA) DNA sequencing. Number in each analysis indicates how many specimens were utilised in a given method (a–adults, j–juveniles, l–larvae)." httpUri="http://table.plazi.org/id/5DCF37ADB944FFF21CEDFE7FBF58FDBB" pageId="39" pageNumber="40" tableUuid="5DCF37ADB944FFF21CEDFE7FBF58FDBB">Table 1</tableCitation>
for all sequenced populations. Comparative material embraced numerous populations of
<taxonomicName id="CEB01CA6B966FFD01D44F894BB76F8FB" authorityName="Richters" authorityYear="1902" box="[318,519,1851,1877]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="39" pageNumber="40" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B966FFD01D44F894BB76F8FB" box="[318,519,1851,1877]" italics="true" pageId="39" pageNumber="40">E. quadrispinosus</emphasis>
</taxonomicName>
from
<collectingCountry id="71A727B5B966FFD01E30F893BBE8F8FB" box="[586,665,1851,1877]" name="France" pageId="39" pageNumber="40">France</collectingCountry>
,
<collectingCountry id="71A727B5B966FFD01ED9F893BA7DF8FB" box="[675,780,1851,1877]" name="Germany" pageId="39" pageNumber="40">Germany</collectingCountry>
,
<collectingCountry id="71A727B5B966FFD01F6DF893BA3AF8FB" box="[791,843,1851,1877]" name="Italy" pageId="39" pageNumber="40">Italy</collectingCountry>
,
<collectingCountry id="71A727B5B966FFD01F2CF893BAD5F8FB" box="[854,932,1851,1877]" name="Russia" pageId="39" pageNumber="40">Russia</collectingCountry>
,
<collectingCountry id="71A727B5B966FFD01FD4F893BD62F8FB" box="[942,1043,1851,1877]" name="United Kingdom" pageId="39" pageNumber="40">Scotland</collectingCountry>
, and
<collectingCountry id="71A727B5B966FFD01836F893BDFEF8FB" box="[1100,1167,1851,1877]" name="Spain" pageId="39" pageNumber="40">Spain</collectingCountry>
, which has already been recorded from all these countries (
<bibRefCitation id="6D211AD4B966FFD01E6CF8F7BBCCF8D7" author="McInnes, S. J." box="[534,701,1887,1913]" pageId="39" pageNumber="40" pagination="257 - 352" refId="ref35021" refString="McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28 (2), 257 - 352. https: // doi. org / 10.1080 / 00222939400770131" type="journal article" year="1994">McInnes 1994</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="41AA34AEB966FFD01CBDF82ABB5EF86E" pageId="39" pageNumber="40" type="etymology">
<paragraph id="090F6725B966FFD01CBDF82ABB5EF86E" blockId="39.[151,1436,1708,1984]" pageId="39" pageNumber="40">
Etymology: From Latin
<emphasis id="3BC4BB37B966FFD01DAFF82BBB44F832" box="[469,565,1923,1948]" italics="true" pageId="39" pageNumber="40">quattuor</emphasis>
= four +
<emphasis id="3BC4BB37B966FFD01EE1F82BBB8AF832" box="[667,763,1923,1948]" italics="true" pageId="39" pageNumber="40">spinosus</emphasis>
= spiny, underlining the presence of dorsal spines
<emphasis id="3BC4BB37B966FFD01950F82BBC35F83E" box="[1322,1348,1922,1949]" italics="true" pageId="39" pageNumber="40">
<superScript id="FEC5CA6DB966FFD01950F82BBC35F83E" attach="left" box="[1322,1348,1922,1949]" fontSize="6" pageId="39" pageNumber="40">Cd</superScript>
</emphasis>
and
<emphasis id="3BC4BB37B966FFD01903F82BBCE4F83E" box="[1401,1429,1922,1948]" italics="true" pageId="39" pageNumber="40">
<superScript id="FEC5CA6DB966FFD01903F82BBCE4F83E" attach="left" box="[1401,1429,1922,1948]" fontSize="6" pageId="39" pageNumber="40">Dd</superScript>
</emphasis>
. An adjective in nominative singular.
</paragraph>
</subSubSection>
<caption id="5DCF37ADB969FFDF1CE2F83FBBFAF87A" ID-DOI="http://doi.org/10.5281/zenodo.8346103" ID-Zenodo-Dep="8346103" httpUri="https://zenodo.org/record/8346103/files/figure.png" pageId="40" pageNumber="41" startId="40.[152,255,1943,1967]" targetBox="[332,1255,181,1919]" targetPageId="40" targetType="figure">
<paragraph id="090F6725B969FFDF1CE2F83FBBFAF87A" blockId="40.[151,1437,1943,2004]" pageId="40" pageNumber="41">
<emphasis id="3BC4BB37B969FFDF1CE2F83FB859F801" bold="true" box="[152,296,1943,1967]" pageId="40" pageNumber="41">FIGURE 23.</emphasis>
<taxonomicName id="CEB01CA6B969FFDF1D4BF83FBB33F801" authorityName="Richters" authorityYear="1902" box="[305,578,1943,1967]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="40" pageNumber="41" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B969FFDF1D4BF83FBB33F801" box="[305,578,1943,1967]" italics="true" pageId="40" pageNumber="41">Echiniscus quadrispinosus</emphasis>
</taxonomicName>
from Germany (PCM): A—male in dorsolateral view, B—juvenile in dorsolateral view, C—larva in dorsal view. Scale bars = 50 μm.
</paragraph>
</caption>
<caption id="5DCF37ADB968FFDE1CE2FAB6BA6BFAD1" ID-DOI="http://doi.org/10.5281/zenodo.8346105" ID-Zenodo-Dep="8346105" httpUri="https://zenodo.org/record/8346105/files/figure.png" pageId="41" pageNumber="42" startId="41.[152,255,1310,1334]" targetBox="[151,1435,181,1286]" targetPageId="41" targetType="figure">
<paragraph id="090F6725B968FFDE1CE2FAB6BA6BFAD1" blockId="41.[151,1437,1310,1407]" pageId="41" pageNumber="42">
<emphasis id="3BC4BB37B968FFDE1CE2FAB6B852FA98" bold="true" box="[152,291,1310,1334]" pageId="41" pageNumber="42">FIGURE 24.</emphasis>
<taxonomicName id="CEB01CA6B968FFDE1D5DFAB6BB45FA98" authorityName="Richters" authorityYear="1902" box="[295,564,1310,1334]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B968FFDE1D5DFAB6BB45FA98" box="[295,564,1310,1334]" italics="true" pageId="41" pageNumber="42">Echiniscus quadrispinosus</emphasis>
</taxonomicName>
from Germany—cephalic region:A—scapular plate sculpturing in upper magnification (PCM), B—scapular plate sculpturing in lower magnification (PCM), C—scapular plate sculpturing (SEM), D—cephalic appendages and subcephalic plates (SEM). Scale bars = 20 μm.
</paragraph>
</caption>
<subSubSection id="41AA34AEB968FFDE1CBDFA02BA96F952" pageId="41" pageNumber="42" type="description">
<paragraph id="090F6725B968FFDE1CBDFA02BA96F952" blockId="41.[151,1437,1450,2034]" pageId="41" pageNumber="42">
Shortened description. Medium-sized (
<emphasis id="3BC4BB37B968FFDE1EFDFA03BBD0FA6A" box="[647,673,1451,1476]" italics="true" pageId="41" pageNumber="42">ca</emphasis>
. 190–310 μm). Body appendage configuration
<emphasis id="3BC4BB37B968FFDE18B6FA02BCF3FA6A" box="[1228,1410,1450,1476]" italics="true" pageId="41" pageNumber="42">
<superScript id="FEC5CA6DB968FFDE18B6FA02BCF3FA6A" attach="left" box="[1228,1410,1450,1476]" fontSize="6" pageId="41" pageNumber="42">A-B-C-Cd- D-Dd-</superScript>
</emphasis>
E, usually stable and with rare asymmetries. Dorsal appendages always spiniform, whereas lateral ones developed as long filamentous cirri (
<figureCitation id="918B7BA0B968FFDE1DBEFA47BB2EF9A7" box="[452,607,1519,1545]" captionStart-0="FIGURE 22" captionStart-1="FIGURE 23" captionStartId-0="39.[152,255,986,1010]" captionStartId-1="40.[152,255,1943,1967]" captionTargetBox-0="[151,1435,166,970]" captionTargetBox-1="[332,1255,181,1919]" captionTargetId-0="figure-19@39.[151,1435,166,970]" captionTargetId-1="figure-26@40.[332,1255,181,1919]" captionTargetPageId-0="39" captionTargetPageId-1="40" captionText-0="FIGURE 22. Echiniscus quadrispinosus from Germany (females): A—dorsal view (PCM), B—dorsal view (SEM), C— dorsolateral view (PCM), D—lateral view (SEM). Scale bars = 50 μm." captionText-1="FIGURE 23. Echiniscus quadrispinosus from Germany (PCM): A—male in dorsolateral view, B—juvenile in dorsolateral view, C—larva in dorsal view. Scale bars = 50 μm." figureDoi-0="http://doi.org/10.5281/zenodo.8346101" figureDoi-1="http://doi.org/10.5281/zenodo.8346103" httpUri-0="https://zenodo.org/record/8346101/files/figure.png" httpUri-1="https://zenodo.org/record/8346103/files/figure.png" pageId="41" pageNumber="42">Figs 22–23A</figureCitation>
). Dorsal plate sculpturing includes (1) dominant large pores (
<figureCitation id="918B7BA0B968FFDE1953FA47B9DAF982" captionStart="FIGURE 24" captionStartId="41.[152,255,1310,1334]" captionTargetBox="[151,1435,181,1286]" captionTargetId="figure-19@41.[151,1435,181,1286]" captionTargetPageId="41" captionText="FIGURE 24. Echiniscus quadrispinosus from Germany—cephalic region:A—scapular plate sculpturing in upper magnification (PCM), B—scapular plate sculpturing in lower magnification (PCM), C—scapular plate sculpturing (SEM), D—cephalic appendages and subcephalic plates (SEM). Scale bars = 20 μm." figureDoi="http://doi.org/10.5281/zenodo.8346105" httpUri="https://zenodo.org/record/8346105/files/figure.png" pageId="41" pageNumber="42">Figs 24A, C</figureCitation>
) overlapping with (2) densely arranged endocuticular pillars (
<figureCitation id="918B7BA0B968FFDE1F0FF9BABAAEF982" box="[885,991,1554,1580]" captionStart="FIGURE 24" captionStartId="41.[152,255,1310,1334]" captionTargetBox="[151,1435,181,1286]" captionTargetId="figure-19@41.[151,1435,181,1286]" captionTargetPageId="41" captionText="FIGURE 24. Echiniscus quadrispinosus from Germany—cephalic region:A—scapular plate sculpturing in upper magnification (PCM), B—scapular plate sculpturing in lower magnification (PCM), C—scapular plate sculpturing (SEM), D—cephalic appendages and subcephalic plates (SEM). Scale bars = 20 μm." figureDoi="http://doi.org/10.5281/zenodo.8346105" httpUri="https://zenodo.org/record/8346105/files/figure.png" pageId="41" pageNumber="42">Fig. 24B</figureCitation>
). Caudal plate often with unsculptured epicuticular ridges forming a cross. Subcephalic (
<figureCitation id="918B7BA0B968FFDE1EB5F99CBA49F9E1" box="[719,824,1588,1615]" captionStart="FIGURE 24" captionStartId="41.[152,255,1310,1334]" captionTargetBox="[151,1435,181,1286]" captionTargetId="figure-19@41.[151,1435,181,1286]" captionTargetPageId="41" captionText="FIGURE 24. Echiniscus quadrispinosus from Germany—cephalic region:A—scapular plate sculpturing in upper magnification (PCM), B—scapular plate sculpturing in lower magnification (PCM), C—scapular plate sculpturing (SEM), D—cephalic appendages and subcephalic plates (SEM). Scale bars = 20 μm." figureDoi="http://doi.org/10.5281/zenodo.8346105" httpUri="https://zenodo.org/record/8346105/files/figure.png" pageId="41" pageNumber="42">Fig. 24D</figureCitation>
) and genital (
<figureCitation id="918B7BA0B968FFDE1FA2F99CBD31F9E1" box="[984,1088,1588,1615]" captionStart="FIGURE 25" captionStartId="42.[152,255,1345,1369]" captionTargetBox="[151,1435,222,1330]" captionTargetId="figure-46@42.[151,1435,222,1330]" captionTargetPageId="42" captionText="FIGURE 25. Echiniscus quadrispinosus from Germany—morphological details: A—claws III (PCM), B —claws III (SEM), C—claws IV (SEM), D—genital plates and gonopore (SEM). Scale bars = 10 μm." figureDoi="http://doi.org/10.5281/zenodo.8346107" httpUri="https://zenodo.org/record/8346107/files/figure.png" pageId="41" pageNumber="42">Fig. 25D</figureCitation>
) plates present. Dentate collar IV with numerous, long irregular teeth. Claws of a moderate size with respect to the body length, with small primary spurs closely positioned to claw bases (
<figureCitation id="918B7BA0B968FFDE1EC2F9D2BA39F93B" box="[696,840,1658,1685]" captionStart="FIGURE 25" captionStartId="42.[152,255,1345,1369]" captionTargetBox="[151,1435,222,1330]" captionTargetId="figure-46@42.[151,1435,222,1330]" captionTargetPageId="42" captionText="FIGURE 25. Echiniscus quadrispinosus from Germany—morphological details: A—claws III (PCM), B —claws III (SEM), C—claws IV (SEM), D—genital plates and gonopore (SEM). Scale bars = 10 μm." figureDoi="http://doi.org/10.5281/zenodo.8346107" httpUri="https://zenodo.org/record/8346107/files/figure.png" pageId="41" pageNumber="42">Figs 25A–C</figureCitation>
). Juveniles with somewhat fainter dorsal sculpturing and body appendage formula
<emphasis id="3BC4BB37B968FFDE1D9DF935BBF2F918" box="[487,643,1692,1719]" italics="true" pageId="41" pageNumber="42">
<superScript id="FEC5CA6DB968FFDE1D9DF935BBF2F918" attach="left" box="[487,643,1692,1719]" fontSize="6" pageId="41" pageNumber="42">A-C-Cd- D-Dd-</superScript>
</emphasis>
E; clearly smaller than sexually mature individuals (
<figureCitation id="918B7BA0B968FFDE18A9F934BC4AF919" box="[1235,1339,1692,1719]" captionStart="FIGURE 23" captionStartId="40.[152,255,1943,1967]" captionTargetBox="[332,1255,181,1919]" captionTargetId="figure-26@40.[332,1255,181,1919]" captionTargetPageId="40" captionText="FIGURE 23. Echiniscus quadrispinosus from Germany (PCM): A—male in dorsolateral view, B—juvenile in dorsolateral view, C—larva in dorsal view. Scale bars = 50 μm." figureDoi="http://doi.org/10.5281/zenodo.8346103" httpUri="https://zenodo.org/record/8346103/files/figure.png" pageId="41" pageNumber="42">Fig. 23B</figureCitation>
). Larvae much smaller than juveniles, with a pronounced layer of endocuticular pillars and large pores arranged mainly at the posterior margins of plates; body appendage formula
<emphasis id="3BC4BB37B968FFDE1E94F94ABA21F952" box="[750,848,1762,1788]" italics="true" pageId="41" pageNumber="42">
<superScript id="FEC5CA6DB968FFDE1E94F94ABA21F952" attach="left" box="[750,848,1762,1788]" fontSize="6" pageId="41" pageNumber="42">A-Cd- Dd-</superScript>
</emphasis>
E (
<figureCitation id="918B7BA0B968FFDE1F09F94ABAABF952" box="[883,986,1762,1788]" captionStart="FIGURE 23" captionStartId="40.[152,255,1943,1967]" captionTargetBox="[332,1255,181,1919]" captionTargetId="figure-26@40.[332,1255,181,1919]" captionTargetPageId="40" captionText="FIGURE 23. Echiniscus quadrispinosus from Germany (PCM): A—male in dorsolateral view, B—juvenile in dorsolateral view, C—larva in dorsal view. Scale bars = 50 μm." figureDoi="http://doi.org/10.5281/zenodo.8346103" httpUri="https://zenodo.org/record/8346103/files/figure.png" pageId="41" pageNumber="42">Fig. 23C</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="41AA34AEB968FFDD1CBDF8ACB8FCFF7B" lastPageId="42" lastPageNumber="43" pageId="41" pageNumber="42" type="discussion">
<paragraph id="090F6725B968FFDE1CBDF8ACBCEAF8EC" blockId="41.[151,1437,1450,2034]" pageId="41" pageNumber="42">
Phylogenetic position:
<taxonomicName id="CEB01CA6B968FFDE1DB9F8ACBB99F8B0" authorityName="Richters" authorityYear="1902" box="[451,744,1796,1822]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B968FFDE1DB9F8ACBB99F8B0" box="[451,744,1796,1822]" italics="true" pageId="41" pageNumber="42">Echiniscus quadrispinosus</emphasis>
</taxonomicName>
is a sister species of the clade
<taxonomicName id="CEB01CA6B968FFDE1848F8ADBDB6F8B0" box="[1074,1223,1796,1822]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="granulatus">
<emphasis id="3BC4BB37B968FFDE1848F8ADBDB6F8B0" box="[1074,1223,1796,1822]" italics="true" pageId="41" pageNumber="42">E. granulatus</emphasis>
</taxonomicName>
+
<taxonomicName id="CEB01CA6B968FFDE1899F8ADBC2FF8B0" box="[1251,1374,1796,1822]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="militaris">
<emphasis id="3BC4BB37B968FFDE1899F8ADBC2FF8B0" box="[1251,1374,1796,1822]" italics="true" pageId="41" pageNumber="42">E. militaris</emphasis>
</taxonomicName>
(Figs 1–2). The genetic diversity between sequenced populations was low and evinced in a shallow phylogenetic structure.
</paragraph>
<paragraph id="090F6725B968FFDD1CBDF8E2B8FCFF7B" blockId="41.[151,1437,1450,2034]" lastBlockId="42.[151,1436,151,213]" lastPageId="42" lastPageNumber="43" pageId="41" pageNumber="42">
Remarks: The biogeographic range of
<taxonomicName id="CEB01CA6B968FFDE1E09F8E3BA4CF8CA" authorityName="Richters" authorityYear="1902" box="[627,829,1866,1892]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B968FFDE1E09F8E3BA4CF8CA" box="[627,829,1866,1892]" italics="true" pageId="41" pageNumber="42">E. quadrispinosus</emphasis>
</taxonomicName>
is difficult to assess due to amassed misidentifications throughout the globe (
<bibRefCitation id="6D211AD4B968FFDE1DE8F8C5BB4BF829" author="McInnes, S. J." box="[402,570,1901,1927]" pageId="41" pageNumber="42" pagination="257 - 352" refId="ref35021" refString="McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28 (2), 257 - 352. https: // doi. org / 10.1080 / 00222939400770131" type="journal article" year="1994">McInnes 1994</bibRefCitation>
). The well-documented and trustworthy records of
<taxonomicName id="CEB01CA6B968FFDE1804F8C5BC38F828" authorityName="Richters" authorityYear="1902" box="[1150,1353,1900,1926]" class="Heterotardigrada" family="Echiniscidae" genus="Echiniscus" kingdom="Animalia" order="Echiniscoidea" pageId="41" pageNumber="42" phylum="Tardigrada" rank="species" species="quadrispinosus">
<emphasis id="3BC4BB37B968FFDE1804F8C5BC38F828" box="[1150,1353,1900,1926]" italics="true" pageId="41" pageNumber="42">E. quadrispinosus</emphasis>
</taxonomicName>
extend from Macaronesia and Spain in Western Palaearctic to the Caucasus in Central Palaearctic (
<bibRefCitation id="6D211AD4B968FFDE189AF827B8B1F863" author="da Cunha, A. X. &amp; do Nascimento Ribeiro, F." pageId="41" pageNumber="42" pagination="1 - 24" refId="ref29935" refString="da Cunha, A. X. &amp; do Nascimento Ribeiro, F. (1962) A fauna de tardigrados da Ilha da Madeira. Memorias e Estudos do Museu Zoologico da Universidade de Coimbra, 279, 1 - 24." type="journal article" year="1962">
da
<collectingCountry id="71A727B5B968FFDE1972F827BC20F807" box="[1288,1361,1935,1961]" name="Saint Helena, Ascension and Tristan da Cunha" pageId="41" pageNumber="42">Cunha</collectingCountry>
&amp; do Nascimento Ribeiro 1962
</bibRefCitation>
;
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;
<bibRefCitation id="6D211AD4B968FFDE1E00F81BBB83F863" author="Guil, N." box="[634,754,1971,1997]" pageId="41" pageNumber="42" pagination="75 - 94" refId="ref32636" refString="Guil, N. (2002) Diversity and distribution of tardigrades (Bilateria, Tardigrada) from the Iberian Peninsula, Balearic Islands and Chafarinas Islands. Graellsia, 58 (2), 75 - 94. https: // doi. org / 10.3989 / graellsia. 2002. v 58. i 2.279" type="journal article" year="2002">Guil 2002</bibRefCitation>
; present study). We do not exclude the possibility that this species inhabits Eastern Palaearctic and the Nearctic region, making it a Holarctic element, but the current data do not allow for claiming that with certainty. Reports from other zoogeographic regions are highly dubious as based mostly on chaetotaxy.
</paragraph>
</subSubSection>
</treatment>
</document>
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