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<mods:title id="E0B0E999EDE6B9ECC6FA856F1438FBAE">Taxonomy of North European Lumbricillus (Clitellata, Enchytraeidae)</mods:title>
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<mods:namePart id="94CBBEB386CAB5F0105EC9EE9FA6B36A">Klinth, Marten J.</mods:namePart>
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<mods:namePart id="592D3BD337B79CD5C1F22D158D23F9DA">Rota, Emilia</mods:namePart>
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<mods:namePart id="78C16322BA1B2F8717D1EA291739E739">Erseus, Christer</mods:namePart>
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<treatment id="4F4397069F4BD930D24731E5F4A501AE" ID-GBIF-Taxon="134860661" LSID="urn:lsid:plazi:treatment:4F4397069F4BD930D24731E5F4A501AE" httpUri="http://treatment.plazi.org/id/4F4397069F4BD930D24731E5F4A501AE" lastPageNumber="47" pageId="32" pageNumber="47">
<subSubSection id="A460B5A50E918BDA06C1F00601F7A110" pageId="32" pageNumber="47" type="nomenclature">
<paragraph id="94B5A02797E3DF8441256F9A3F94D80A" pageId="32" pageNumber="47">
<taxonomicName id="68057E935451206D8B693EFDF90EE5C4" ID-CoL="6R2DS" authority="Michaelsen, 1889" authorityName="Michaelsen" authorityYear="1889" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">
<pageBreakToken id="AD3FE3F4193AAE3D8DB500167ADC4BF6" pageId="32" pageNumber="47" start="start">Lumbricillus</pageBreakToken>
arenarius (Michaelsen, 1889)
</taxonomicName>
Fig. 21
</paragraph>
</subSubSection>
<subSubSection id="D60F822C1ED8FE0BC0C6EC0924273F10" pageId="32" pageNumber="47" type="reference_group">
<paragraph id="DEEE8FF58F93CA95D171B49803E672E1" pageId="32" pageNumber="47">
<taxonomicName id="1F30C3684B47DF2DD202110DCEC5D549" class="Clitellata" family="Enchytraeidae" genus="Enchytraeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Enchytraeus arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">Enchytraeus arenarius</taxonomicName>
Michaelsen, 1889: pp. 12-14, figs 5
<normalizedToken id="257B8CE011E7B4EB01184C131E0ED846" originalValue="a–d">a-d</normalizedToken>
.
</paragraph>
<paragraph id="3960FB2095F39041903673BF18A5B646" pageId="32" pageNumber="47">
<taxonomicName id="8F5E017E489DA7A084C86E97CD04D3A5" class="Clitellata" family="Enchytraeidae" genus="Marionina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Marionina arenaria" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenaria">Marionina arenaria</taxonomicName>
;
<bibRefCitation id="D9BA6BDCA4EC6C35FFC080323FA73C35" author="Michaelsen, W" journalOrPublisher="Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten" pageId="47" pageNumber="62" url="https://doi.org/10.5962/bhl" year="1900">Michaelsen 1900</bibRefCitation>
: pp. 74-75.
</paragraph>
<paragraph id="F4D2CB735A14EA9CE8E396EAD50EFBC8" pageId="32" pageNumber="47">
<taxonomicName id="698834B19E034A2E481126C58AB77D0A" class="Clitellata" family="Enchytraeidae" genus="Enchytraeoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Enchytraeoides arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">Enchytraeoides arenarius</taxonomicName>
;
<bibRefCitation id="77BC6071A6AB044437348C1C3B9DAA3B" author="Ude, H" journalOrPublisher="Die Tierwelt Deutschlands" pageId="50" pageNumber="65" pagination="1 - 132" title="Oligochaeta." volume="15" year="1929">Ude 1929</bibRefCitation>
: pp. 62-63;
<bibRefCitation id="A159EABCFBD2360543A86CC834D7A11F" author="von Buelow, T" journalOrPublisher="Kieler Meeresforschungen" pageId="44" pageNumber="59" pagination="69 - 116" title="Systematisch-autoekologische Studien an eulitoralen Oligochaeten der Kimbrischen Halbinsel." volume="13" year="1957">
von
<normalizedToken id="8B7C734F688CC98A8BAA6BDD7F6A262E" originalValue="Bülow">Buelow</normalizedToken>
1957
</bibRefCitation>
: p. 84;
<bibRefCitation id="8DF5B9B87FB1F10072C3B01A435E6F63" author="Knoellner, FH" journalOrPublisher="Abteilung fuer Systematik, Geographie und Biologie der Tier" pageId="46" pageNumber="61" pagination="425 - 563" title="Oekologische und systematische Untersuchungen ueber litorale und marine Oligochaeten der Kieler Bucht. Zoologische Jahrbuecher." volume="66" year="1935">
<normalizedToken id="29D9D51F01B07B554183D2B9502417F6" originalValue="Knöllner">Knoellner</normalizedToken>
1935
</bibRefCitation>
: pp. 437-438, figs 7-8.
</paragraph>
<paragraph id="C84321C7117EF80F23081B71F3391D78" pageId="32" pageNumber="47">
<taxonomicName id="4F91ED22ACABEFDB4429974BB57D44B6" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">Lumbricillus arenarius</taxonomicName>
;
<bibRefCitation id="830346AA6992D3F5FFECC01F7028872F" pageId="32" pageNumber="47">Nielsen and Christensen 1959</bibRefCitation>
: pp. 107-108, figs 127-128;
<bibRefCitation id="414DD3EB63E7D40985FD58DA81C60E56" author="Rota, E" journalOrPublisher="Journal of Natural History" pageId="48" pageNumber="63" pagination="29 - 64" title="A taxonomic study of some Swedish Enchytraeidae (Oligochaeta), with descriptions of four new species and notes on the genus Fridericia." url="https://doi.org/10.1080/002229399300461" volume="33" year="1999">Rota and Healy 1999</bibRefCitation>
: pp. 53-54;
<bibRefCitation id="833ECECADDADE74DC0217FE24E916B3A" author="Erseus, C" journalOrPublisher="Hydrobiologia" pageId="45" pageNumber="60" pagination="101 - 110" title="Clitellate diversity in Nationalstadsparken, an urban national park in Stockholm, Sweden." url="https://doi.org/10.1007/978-94-011-4207-6_10" volume="406" year="1999">
<normalizedToken id="7AD10A6C2E4D3C51A2A57283E775471B" originalValue="Erséus">Erseus</normalizedToken>
et al. 1999
</bibRefCitation>
;
<bibRefCitation id="A9A26C423084ACBE9BE53EF14E334685" author="Erseus, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="46" pageNumber="61" pagination="849 - 858" title="Molecular phylogeny of Enchytraeidae (Annelida, Clitellata)." url="https://doi.org/10.1016/j.ympev.2010.07.005" volume="57" year="2010">
<normalizedToken id="92525157E49EDFF65F31C768F425189C" originalValue="Erséus">Erseus</normalizedToken>
et al. 2010
</bibRefCitation>
;
<bibRefCitation id="794DE10623B1569D0C631E5A549E18B9" author="Klinth, MJ" journalOrPublisher="Zoologica Scripta" pageId="46" pageNumber="61" pagination="96 - 110" title="Phylogeny and species delimitation of North European Lumbricillus (Clitellata, Enchytraeidae)." url="https://doi.org/10.1111/zsc.12187" volume="46" year="2017">Klinth et al. 2017</bibRefCitation>
.
</paragraph>
<paragraph id="5DAB35319FF5CF175AF718B1D4253BE6" pageId="32" pageNumber="47">
<taxonomicName id="A9CFA41B462B874442F0FB0C16DD1044" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus magdalenae" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="magdalenae">Lumbricillus magdalenae</taxonomicName>
Nurminen, 1965: pp. 6-7, figs 2
<normalizedToken id="81E2DCDE383B4621AF82381C3368C07A" originalValue="e–g">e-g</normalizedToken>
.
</paragraph>
</subSubSection>
<subSubSection id="FF6FB799B27A03308FAA56817F14481A" pageId="32" pageNumber="47" type="type material">
<paragraph id="48EA397E90D417343BAB4F91B0B494C2" pageId="32" pageNumber="47">Type material.</paragraph>
<paragraph id="D7F249392D7E2B52117A043DFE4B78D4" pageId="32" pageNumber="47">
Typus amissus (Nomenclatura Oligochaetologica). Type locality: Elbe River, Hamburg, Germany (
<bibRefCitation id="1EDF37CFFD6F597B94B0F853D9DC99A9" author="Michaelsen, W" journalOrPublisher="Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten" pageId="47" pageNumber="62" pagination="1 - 17" title="Oligochaeten des Naturhistorischen Museums in Hamburg." volume="6" year="1889">Michaelsen 1889</bibRefCitation>
). We did not designate a neotype as we do not have material from the type locality.
</paragraph>
</subSubSection>
<subSubSection id="099E90434D5DE2A1253699FFDE1F87D1" pageId="32" pageNumber="47" type="materials_examined">
<paragraph id="F6E13C80F27CF4DCB711EB3145FF9EC9" pageId="32" pageNumber="47">Material examined.</paragraph>
<paragraph id="A0D50F79DEFC84F0663F90A97C5A8096" pageId="32" pageNumber="47">SMNH 152716 (CE1001), one mature specimen from Sweden, and ZMBN 107784 (CE8474), ZMBN 107787 (CE20748), ZMBN 107788 (CE20749) &amp; ZMBN 107789 (CE20750), four mature specimens from Norway. For information on specimen collection localities and GenBank accession numbers see Appendix 1.</paragraph>
</subSubSection>
<subSubSection id="DE5A45621349DC41EF90F7A25D08EE46" pageId="32" pageNumber="47" type="description">
<paragraph id="65C4CB0FA511BC1B578683270021FE52" pageId="32" pageNumber="47">Description.</paragraph>
<paragraph id="FA195D79D513376F4839F2062888F1E4" pageId="32" pageNumber="47">
White to yellow worms. Length (fixed worms) more than 5.0-8.6 mm (amputated specimens), first 15 segments 3.5-4.0 mm long, width at clitellum 0.31-0.51 mm. More than 19-35 segments. Chaetae straight or slightly sigmoid (Fig. 21A). Lateral bundles with 2-3 chaetae anterior to clitellum, 2 chaetae in postclitellar segments. Ventral bundles with 2-3(4) chaetae anterior to clitellum, 2-3 chaetae posteriorly. Each
<normalizedToken id="AAC54ACE67AC3FA14D467A30FA8E3669" originalValue="worm’s">worm's</normalizedToken>
longest measured chaetae 40-70
<normalizedToken id="F78B8BDDD40A2F9FA5ACF7E0624796A7" originalValue="µm">µm</normalizedToken>
long, about 5
<normalizedToken id="4D729ECBD30A380013CC14E63D7438B6" originalValue="µm">µm</normalizedToken>
wide. Clitellum extending over
<normalizedToken id="DAB070EB43A6AD4BD1D6E40E0A5DE2E3" originalValue="XII–">XII-</normalizedToken>
1/2XIII, in some covering all of XIII. Head pore at 0/1. Epidermis with transverse rows of gland cells.
</paragraph>
<paragraph id="ABD8D2676CACA3E8EF3F6E6DAEC02A03" pageId="32" pageNumber="47">
Coelomocytes numerous, 20-50
<normalizedToken id="C73666BC0BDF52AA74BA32E644D0F227" originalValue="µm">µm</normalizedToken>
long, spindle-shaped, oval, round, granulated with distinct nucleus, some with distally hooked ends. Paired pharyngeal glands present in IV, V and VI (Fig. 21B). Dorsal vessel originating in XIII. Nephridia observed in
<normalizedToken id="033EEE8F72478637AEE120EEF1BFE707" originalValue="XV–XVI">XV-XVI</normalizedToken>
and
<normalizedToken id="54210C05F137FE6D9552795DBE8D9408" originalValue="XX–XXV">XX-XXV</normalizedToken>
, 105-145
<normalizedToken id="0CE124314CDBF0D44B4CB636053DA7CB" originalValue="µm">µm</normalizedToken>
long, anteseptale funnel only, postseptale oval, tapering into efferent duct. Brain with posterior incision.
</paragraph>
<paragraph id="16441B3ADC25436A6495F82680FFFDA1" pageId="32" pageNumber="47">
Male genitalia paired (Fig. 21D). Testes originating in XI, extending forwards into X, with testis sacs covering mass of rather large irregularly arranged lobes. Sperm funnels in XI, in some specimens extending back into XII, 375-975
<normalizedToken id="62CD131E6340018DACBDC4045E186C84" originalValue="µm">µm</normalizedToken>
long, 55-103
<normalizedToken id="3F8B61E46BAD6E9C7CD3F4276DB68A3B" originalValue="µm">µm</normalizedToken>
wide, making them 6-13 times longer than wide, funnels tapering towards vasa deferentia. Most of vasa irregularly coiled in XII, in one specimen extending back into XIV, 5-10
<normalizedToken id="78BC98F44F6023AEE6EE69F0AEC0E972" originalValue="µm">µm</normalizedToken>
wide. Penial bulbs round, 110-140
<normalizedToken id="4B6368BF64C354C20BAF0334A22E9EE9" originalValue="µm">µm</normalizedToken>
in diameter. Ovaries in XII. One to six mature eggs present at a time.
</paragraph>
<paragraph id="C22FD2DBD2D086EFE3921A7C11857191" pageId="32" pageNumber="47">
Spermathecae (Fig. 21C) in V, pouch-shaped. Ectal duct longer than and gradually widening into ampulla. Ampulla oval or round, entally connecting with oesophagus. Irregular mass of sperm aggregated in ampulla. Spermathecae 100-255
<normalizedToken id="5447DFA34051E5DDCFF7259E8C513C92" originalValue="µm">µm</normalizedToken>
long, 50-115
<normalizedToken id="56A00D65D08C4AEF258E05C4FA8D970A" originalValue="µm">µm</normalizedToken>
wide at widest part of ampulla. Gland cells surrounding ectal pore, divided into several flaps, whole glandular body 75-135
<normalizedToken id="294B1A0B144F96F63E087875048C6101" originalValue="µm">µm</normalizedToken>
in diameter at its widest part. Up to four midventral subneural glands in
<normalizedToken id="87FB52EAF6AC41359A3CC1E71D6CCDEC" originalValue="XIII–XVI">XIII-XVI</normalizedToken>
, 75-110
<normalizedToken id="CE229AA7FA91631F31F4EB15513A86EC" originalValue="µm">µm</normalizedToken>
, 90-115
<normalizedToken id="03AD9BC5870968E956ABF9E3DEA95D37" originalValue="µm">µm</normalizedToken>
, 75-85
<normalizedToken id="5045D16B467CA4337BB5996279A50154" originalValue="µm">µm</normalizedToken>
and 95
<normalizedToken id="2156C5A17B8D109E1420B56E407CFCB1" originalValue="µm">µm</normalizedToken>
long, respectively; glands in XVI not observed in all specimens.
</paragraph>
<caption id="174C67D149DF0BD25ED81EBC4C38135C" pageId="32" pageNumber="47">
<paragraph id="850633783B03B475384F2C646C8977FA" pageId="32" pageNumber="47">
Figure 21.
<taxonomicName id="CBDA0F5E7BCE7D4B261A07EF4C931F76" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">Lumbricillus arenarius</taxonomicName>
. A Chaetal bundle B Anterior body C Spermatheca D Other genitalia. Abbreviations under general notes. Scale bars: 100
<normalizedToken id="A35AFA8D28DEC0C00B76C885AAAFA715" originalValue="µm">µm</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="928BB77E9E9FF5D2AFB27253C1B18667" pageId="32" pageNumber="47" type="distribution">
<paragraph id="B00CEF4360C7504EBC77540318161B9B" pageId="32" pageNumber="47">Geographical distribution including BOLD data.</paragraph>
<paragraph id="4264C1639AC2A299E77A21D385766CE7" pageId="32" pageNumber="47">
Genetically identified from Norway (mainland and Svalbard) and Sweden. Also reported from Denmark, Canada, Germany, Greenland, Iceland, Ireland, Wales and
<normalizedToken id="D6AA3A4DEC99D3D8BF36BEABD1DAAE7A" originalValue="North–Western">North-Western</normalizedToken>
Australia (
<bibRefCitation id="BFAF3F03E1667395EFDEC6F3EED938A2" author="Rota, E" journalOrPublisher="Journal of Natural History" pageId="48" pageNumber="63" pagination="29 - 64" title="A taxonomic study of some Swedish Enchytraeidae (Oligochaeta), with descriptions of four new species and notes on the genus Fridericia." url="https://doi.org/10.1080/002229399300461" volume="33" year="1999">Rota and Healy 1999</bibRefCitation>
). BIN-number: BOLD:AAT8953.
</paragraph>
</subSubSection>
<subSubSection id="72F9EEEDDE8926818C56F124F9643E7D" pageId="32" pageNumber="47" type="remarks">
<paragraph id="8DBEB80D568A572CFC3FA8A1A79A6E42" pageId="32" pageNumber="47">Remarks.</paragraph>
<paragraph id="CAE2A313D393DB7F56C4FFBA10EC8477" pageId="32" pageNumber="47">
The original description by
<bibRefCitation id="0C398434EFA88B7920FC0B004C7F55E2" author="Michaelsen, W" journalOrPublisher="Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten" pageId="47" pageNumber="62" pagination="1 - 17" title="Oligochaeten des Naturhistorischen Museums in Hamburg." volume="6" year="1889">Michaelsen (1889)</bibRefCitation>
was later amended by
<bibRefCitation id="9B86DE12766F3083F4169326E83E4AFC" author="Knoellner, FH" journalOrPublisher="Abteilung fuer Systematik, Geographie und Biologie der Tier" pageId="46" pageNumber="61" pagination="425 - 563" title="Oekologische und systematische Untersuchungen ueber litorale und marine Oligochaeten der Kieler Bucht. Zoologische Jahrbuecher." volume="66" year="1935">
<normalizedToken id="319901AF063C265F6111382571AE01B2" originalValue="Knöllner">Knoellner</normalizedToken>
(1935)
</bibRefCitation>
who redrew the shape of the nephridia and spermathecae, also confirmed by
<bibRefCitation id="F8098ECCD7CB3D313C83317D8532CC37" pageId="32" pageNumber="47">Nielsen and Christensen (1959)</bibRefCitation>
. The newly examined material in this study resembles the original description in most characters but the spermathecae and nephridia are in agreement with the amended descriptions. Coelomic corpuscles were found with hooked ends which seemed to bind to the internal tissue in a way that is described by Michaelsen. The testes seemed to be either an irregular compact mass or divided into separate lobes, encased in testis sacs, but these lobes were not arranged in the bunch-shape seen in the
<taxonomicName id="598B1829A42B2E5E5B23C8157216A036" lsidName="lineatus" pageId="32" pageNumber="47" rank="species" species="lineatus">lineatus</taxonomicName>
,
<taxonomicName id="364BA55D98E8159D61F9EC51768E4C6C" lsidName="pagenstecheri" pageId="32" pageNumber="47" rank="species" species="pagenstecheri">pagenstecheri</taxonomicName>
and &quot;
<taxonomicName id="5A44223C1F6D9573148E01C562F2432A" lsidName="tuba" pageId="32" pageNumber="47" rank="species" species="tuba">tuba</taxonomicName>
&quot; groups.
</paragraph>
<paragraph id="4088699BF1B51B944C73EA806A904D60" pageId="32" pageNumber="47">
<taxonomicName id="7D545399812BD03FACA8AD8A0E6C71B7" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus arenarius" order="Enchytraeida" pageId="32" pageNumber="47" phylum="Annelida" rank="species" species="arenarius">Lumbricillus arenarius</taxonomicName>
is genetically closely related to
<taxonomicName id="E8DD7C35A7FC38720BA3C5B772B176B6" genus="L." lsidName="L." pageId="32" pageNumber="47" rank="genus">L.</taxonomicName>
sp. H and
<taxonomicName id="43075661BF26901F2D57A70D4D426BDF" lsidName="L. dubius" pageId="32" pageNumber="47" rank="species" species="dubius">L. dubius</taxonomicName>
(Fig. 1).
</paragraph>
</subSubSection>
</treatment>
</document>
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