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<document id="F4B7D3B90F6E8B85BD37D8926350A3F2" ID-CLB-Dataset="31944" ID-DOI="http://dx.doi.org/10.3897/zookeys.703.13385" ID-GBIF-Dataset="e1e83ca0-ed07-4f17-bd7e-1cedbac15bbc" ID-PMC="PMC5673978" ID-Pensoft-Pub="1313-2970-703-15" ID-PubMed="29118604" ID-ZooBank="9BAAB4A5CDE1493B8A0413D8F301E198" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-703-15" ModsDocOrigin="ZooKeys 703" ModsDocTitle="Taxonomy of North European Lumbricillus (Clitellata, Enchytraeidae)" checkinTime="1506576699266" checkinUser="pensoft" docAuthor="Klinth, Marten J., Rota, Emilia &amp; Erseus, Christer" docDate="2017" docId="A4458E15A0C35C51F6C11A042EF16184" docLanguage="en" docName="ZooKeys 703: 15-96" docOrigin="ZooKeys 703" docSource="http://dx.doi.org/10.3897/zookeys.703.13385" docTitle="Lumbricillus lineatus Mueller 1774" docType="treatment" docVersion="6" lastPageNumber="22" masterDocId="B248FFB4FF91C20DCE56FFE7FF875A5A" masterDocTitle="Taxonomy of North European Lumbricillus (Clitellata, Enchytraeidae)" masterLastPageNumber="96" masterPageNumber="15" pageNumber="20" updateTime="1701378398428" updateUser="ExternalLinkService">
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<mods:title id="7028FD9032F8A5497CD97D7E3758A1E7">Taxonomy of North European Lumbricillus (Clitellata, Enchytraeidae)</mods:title>
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<mods:namePart id="CDE954533CCECC4BD92062AC6B9A8BB8">Klinth, Marten J.</mods:namePart>
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<mods:namePart id="91BFA2C0A327CEAFF6E72964FA7CCB16">Rota, Emilia</mods:namePart>
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<mods:namePart id="16928CCB6C4875749D53C4553D57BA42">Erseus, Christer</mods:namePart>
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<mods:date id="972F3C7C43069B28DB8C7F8B98216829">2017</mods:date>
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<mods:identifier id="E760C6F60DF6D637C3EBDDF4F0406B28" type="Pensoft-Pub">1313-2970-703-15</mods:identifier>
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<treatment id="A4458E15A0C35C51F6C11A042EF16184" ID-GBIF-Taxon="134860656" LSID="urn:lsid:plazi:treatment:A4458E15A0C35C51F6C11A042EF16184" httpUri="http://treatment.plazi.org/id/A4458E15A0C35C51F6C11A042EF16184" lastPageId="7" lastPageNumber="22" pageId="5" pageNumber="20">
<subSubSection id="30E3211D8E541E8945083E24C8B41AD5" pageId="5" pageNumber="20" type="nomenclature">
<paragraph id="8D2B529D062B67378A1ACFC120071AC4" pageId="5" pageNumber="20">
<taxonomicName id="16E841974D460FF3F179CD895A63568A" ID-CoL="72Q5X" authority="Mueller, 1774" authorityName="Mueller" authorityYear="1774" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">
Lumbricillus lineatus (
<normalizedToken id="051C825EE195E93885E595CA3C36107D" originalValue="Müller">Mueller</normalizedToken>
, 1774)
</taxonomicName>
Figs 2, 3A
</paragraph>
</subSubSection>
<subSubSection id="290AA895DC00E66CE1B111AC0335E3CA" pageId="5" pageNumber="20" type="reference_group">
<paragraph id="5A5B35C38DA3795ED35B13145627EDEE" pageId="5" pageNumber="20">
<taxonomicName id="68DA881C008DB7D47EE392294B4AE343" class="Clitellata" family="Enchytraeidae" genus="Lumbricus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricus lineatus</taxonomicName>
<normalizedToken id="2339011C3C7797681A23007CF437EBAA" originalValue="Müller">Mueller</normalizedToken>
, 1774: p. 29.
</paragraph>
<paragraph id="41CBF69C9A240CF0092738F9F9343EFF" pageId="5" pageNumber="20">
<taxonomicName id="3B5DB6CF7A90FA6BA017211FE4DDBDB8" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
;
<bibRefCitation id="586359568C35B59547848EA2521C11F9" 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="18281F3987C2B5FD8682E10121BBA624" originalValue="Erséus">Erseus</normalizedToken>
et al. 1999
</bibRefCitation>
;
<bibRefCitation id="0D4231D4FBACC9095446880DDBC58D0A" 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="0B0AA442862B215097D8ACDFFACE3474" originalValue="Erséus">Erseus</normalizedToken>
et al. 2010
</bibRefCitation>
;
<bibRefCitation id="C0E3A1339D14E1AA6CB3B914B572C245" 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="D52B9A8FC5EE1DC17AA0AC7B4A1849E9" pageId="5" pageNumber="20">
<taxonomicName id="E968C65DF9552A65AB11E921056669B7" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pachydrilus claparedeanus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="claparedeanus">Pachydrilus claparedeanus</taxonomicName>
Ditlevsen, 1904: pp. 431-435, figs 28
<normalizedToken id="97873A3B13566DCA9B4690EC8F739E79" originalValue="a–b">a-b</normalizedToken>
.
</paragraph>
<paragraph id="5B8F6C806C8B71854E8D97F3A613AE7E" pageId="5" pageNumber="20">
<taxonomicName id="CFB3546903A0BF6A264C6F5DA6F63A0E" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus agilis" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="agilis">Lumbricillus agilis</taxonomicName>
Moore, 1905: pp. 395-397, pl. XXXIII, figs 23-28.
</paragraph>
<paragraph id="1F3F2ED48BFA1FDA1CD82D4961FA6155" pageId="5" pageNumber="20">
<taxonomicName id="C3F160C14C0741E057CBFA52B4528C0D" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
partim;
<bibRefCitation id="59231225C1CCABED80478CB47D442DB4" 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>
: p. 80;
<bibRefCitation id="EAE4677B24941475A085B555CEDFE23C" author="Welch, PS" journalOrPublisher="Transactions of the American Microscopical Society" pageId="50" pageNumber="65" pagination="119 - 138" title="The Enchytraeidae (Oligochaeta) of the Woods Hole Region, Mass." url="https://doi.org/10.2307/3221697" volume="36" year="1917">Welch 1917</bibRefCitation>
: pp. 123-130;
<bibRefCitation id="DBE2BA7570CD9EFA97C0DF7DA949D100" pageId="5" pageNumber="20">Nielsen and Christensen 1959</bibRefCitation>
: pp. 100-102, figs 109-112.
</paragraph>
<paragraph id="C2DE1E1854C63E623FF38249291C4484" pageId="5" pageNumber="20">
&quot;
<taxonomicName id="C2E5C20D0C551D8BD0AEDAF210F9FD89" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
L2&quot;; BOLD (unpublished records)
</paragraph>
<paragraph id="9C8A65DD7BAE2FDC987908587125A96F" pageId="5" pageNumber="20">
Non
<taxonomicName id="95342D666CF50D65A92ED188F9F12DAD" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pachydrilus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Pachydrilus lineatus</taxonomicName>
; sensu Backlund 1947: pp. 3-5, figs 1-2 (see
<taxonomicName id="44AB2DA4B9E244E130752C228E97B3BC" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus latithecatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="latithecatus">Lumbricillus latithecatus</taxonomicName>
sp. n. below).
</paragraph>
</subSubSection>
<subSubSection id="2C3AE0161CAB0FC3722BE2ADB6CD21A1" pageId="5" pageNumber="20" type="type material">
<paragraph id="F683204B74FB65A87C55E322871EE098" pageId="5" pageNumber="20">Type material (neotype).</paragraph>
<paragraph id="86CBB37F129BA87894F5A72DCF9B1A5C" pageId="5" pageNumber="20">
<taxonomicName id="45AB8012C55ED23EAF69343CFE96214E" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
was described long before reference to types had become common practice and there is no remaining original material (&quot;Typus amissus&quot; in Nomenclatura Oligochaetologica). We designate SMNH Type-8931 [former SMNH 152751] (CE1894) as a neotype of this species. It is a whole-mounted voucher of a sexually mature and DNA-barcoded worm (COI barcode is KU894040 in NCBI/GenBank;
<bibRefCitation id="02C786679DC847BD673A7365C54F456B" 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>
) from the Baltic Sea,
<normalizedToken id="E8A4DFE8CB49C2EA279BC5E1BCD554FA" originalValue="Öland">Oeland</normalizedToken>
,
<normalizedToken id="80AAB6513EED9912DDEE2008DE755594" originalValue="Mörbylånga">Moerbylanga</normalizedToken>
,
<normalizedToken id="1254C7B597519785C995AED10136D22D" originalValue="Skärlöv">Skaerloev</normalizedToken>
fishing harbor, on beach with mixed shelly sand, pebbles and organic material,
<geoCoordinate id="88879F8C07DC0DDD7D663E63051C8C3C" direction="north" orientation="latitude" precision="5" value="56.4241">56.4241 N</geoCoordinate>
,
<geoCoordinate id="660E668474B6F20E5B1294ABB5ABFFD4" direction="east" orientation="longitude" precision="5" value="16.5815">16.5815 E</geoCoordinate>
, collected 10 June 2006 by Lisa Matamoros, Sweden. Our decision to designate this neotype is further discussed in Remarks below.
</paragraph>
</subSubSection>
<subSubSection id="88099559B07B7ECBE551BFA263325A2F" pageId="5" pageNumber="20" type="materials_examined">
<paragraph id="E79EBDA0628E866B3030CA37846D41F6" pageId="5" pageNumber="20">Other material examined.</paragraph>
<paragraph id="5D37D58ECFD825D60C13A2C536C91654" pageId="5" pageNumber="20">SMNH 152742 (CE1640), one mature specimen from Sweden; SMNH 152744 (CE1694), one mature avesiculate triploid specimen from Spain; ZMBN 107872 (CE12043), ZMBN 107875 (CE21688), ZMBN 107879 (CE22604) &amp; ZMBN 107880 (CE22605), four mature specimens from Norway; ZMBN 107878 (CE21986), one mature avesiculate triploid specimen from Norway. For information on collection localities and GenBank accession numbers for COI barcodes see Appendix 1.</paragraph>
</subSubSection>
<subSubSection id="C0C8C70A1E2209E1828C484350CCC682" pageId="5" pageNumber="20" type="description">
<paragraph id="49B15FD4CB5CA083EF4F344265D63729" pageId="5" pageNumber="20">Description.</paragraph>
<paragraph id="2B6732B2C14F26B152770F8E0793A8D5" pageId="5" pageNumber="20">
Orange, red or pink worms. Length (fixed worms) more than 2.2-5.5 mm (amputated specimens), first 15 segments 2.1-2.8 mm long, width at clitellum 0.45-0.75 mm. More than 14-38 segments. Chaetae sigmoid (Fig. 2A). Upper bundles dorsolateral, with 3-6 chaetae anterior to clitellum, 3-7 chaetae in postclitellar segments. Ventral bundles with 4-8 chaetae anterior to clitellum, 3-7(8) chaetae posteriorly. Each
<normalizedToken id="1560C568DA2E564777060EA946C94191" originalValue="worm’s">worm's</normalizedToken>
longest measured chaetae 50-75
<normalizedToken id="90F809285AAEF544CA554DAF12FAE8C8" originalValue="µm">µm</normalizedToken>
long, about 3-5
<normalizedToken id="C7E72194B67B1E480052706D88E0B404" originalValue="µm">µm</normalizedToken>
wide. Clitellum extending over
<normalizedToken id="96A955E068E5A7C87C99FF9B16074661" originalValue="XII–">XII-</normalizedToken>
1/2XIII/XIII. Head pore not observed.
</paragraph>
<paragraph id="EF9F5BC4FB12279D0CDD0A728DB1D844" pageId="5" pageNumber="20">
Coelomocytes, in some specimens numerous, 10-20
<normalizedToken id="7D1B9A63E26276A592C6F00AF89D7E52" originalValue="µm">µm</normalizedToken>
long, round, oval or spindle-shaped, granulated with distinct nucleus. Paired pharyngeal glands present in IV, V and VI; each pair converging dorsally (Fig. 2B). Dorsal vessel originating in
<normalizedToken id="5B659AB57392DED78FDE8E7E73638636" originalValue="XII–XIV">XII-XIV</normalizedToken>
. Nephridia observed in
<normalizedToken id="DBF4AADA3923C5E889CC76C4A0C56C07" originalValue="VIII–X">VIII-X</normalizedToken>
and
<normalizedToken id="84FF37E40EE4B174DFD7415AF1EAA3EC" originalValue="XIII–XX">XIII-XX</normalizedToken>
, about 85-110
<normalizedToken id="0FEBEB375FB9D10123B91C05B4A0B46B" originalValue="µm">µm</normalizedToken>
long. Anteseptale small, consisting of funnel only. Postseptale oval, tapering into posteroventral efferent duct. Brain with posterior incision.
</paragraph>
<paragraph id="F70338F2B5478E5F8F6409977CE219A3" pageId="5" pageNumber="20">
Male genitalia paired (Fig. 2D). Testes originating in XI, extending forwards into X, sometimes IX, with testis sacs forming regular club-shaped lobes, except in two
<normalizedToken id="CF3BA5F5EF8C0EE631DEEFFEE9EBC302" originalValue="“avesiculate”">&quot;avesiculate&quot;</normalizedToken>
specimens (SMNH 152744 and ZMBN 107878) which have atrophic testes. Sperm funnels in XI, 215-420
<normalizedToken id="B44F5E468A42618F3D169C05C1E803D7" originalValue="µm">µm</normalizedToken>
long, 125-185
<normalizedToken id="03FC9367737C5042E6E81365891807D5" originalValue="µm">µm</normalizedToken>
wide, making them about 1.5-2.5 times longer than wide, funnels tapering towards vasa deferentia. Most of vasa irregularly coiled in XII, 15-20
<normalizedToken id="735FC84CE97E3EC63D1D35F523A01088" originalValue="µm">µm</normalizedToken>
wide. Penial bulbs round, 65-140
<normalizedToken id="18A2B961BDF1907607E3F98AFD1E1D31" originalValue="µm">µm</normalizedToken>
in diameter. Ovaries in XII. One to six mature eggs present at a time.
</paragraph>
<paragraph id="C9636788621BAAF8C6561895E618EFA4" pageId="5" pageNumber="20">
Spermathecae (Figs 2C, 3A) in V, spindle-shaped, without distinct ampulla. Ectal duct very short, widening into ampulla. Ampulla with constriction midway dividing it into sections, ectal part narrow, ental part wider, sometimes circular, connecting with oesophagus. Sperm in lumen of ectal part of ampulla, heads of spermatozoa embedded in wall of ental part of ampulla, forming aggregates. Spermathecae 220-275
<normalizedToken id="74422EAF3593A1A57E04FD501BD46E4F" originalValue="µm">µm</normalizedToken>
long, 60-125
<normalizedToken id="2132E0AA0A4AF624810D2D3263A5F239" originalValue="µm">µm</normalizedToken>
wide at widest part of ampulla. Gland cells surrounding ectal duct at pore, forming compact mass, glandular body 80-150
<normalizedToken id="F69570951228DD2906788A841429F055" originalValue="µm">µm</normalizedToken>
in diameter at its widest part. Two midventral subneural glands in
<normalizedToken id="433FD1EA3A6EAF512C49DEC47119E4BE" originalValue="XIII–XIV">XIII-XIV</normalizedToken>
, 60-110
<normalizedToken id="CFECB71CBEB8FA1886F73AA9D23DCA36" originalValue="µm">µm</normalizedToken>
and 60-95
<normalizedToken id="563194A2F3AD0BE51530CC6979D8396F" originalValue="µm">µm</normalizedToken>
long, respectively.
</paragraph>
<caption id="1E7497F1DA03CBC749BCD320570BDF7E" pageId="5" pageNumber="20">
<paragraph id="C809089782474C519BC1B5D5350282D1" pageId="5" pageNumber="20">
Figure 2.
<taxonomicName id="E6D9B8E8624C3A80FE1487B79ADF186F" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
. A Chaetal bundle B Anterior body C Spermatheca (neotype) D Other genitalia. Abbreviations under general notes. Scale bars: 100
<normalizedToken id="B2A340ADD6F6955A1398D09C12042128" originalValue="µm">µm</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="674188933B0F79D2D9F636DF0345CE3C" pageId="5" pageNumber="20" type="details of neotype">
<paragraph id="C3021B81C655E9362E0F3E3E0C4B69FB" pageId="5" pageNumber="20">Details of neotype.</paragraph>
<paragraph id="9B4D6BD16ECAF2F979473996D60BDB73" pageId="5" pageNumber="20">
Length 3.3 mm (amputated specimen), first 15 segments 2.4 mm long, width at clitellum 0.45 mm. 20 segments. Dorsal bundles with 4-6 chaetae anterior to clitellum, 3-4 chaetae in postclitellar segments. Ventral bundles with 6-8 chaetae anterior to clitellum, 3-5 chaetae posteriorly. Longest chaetae 50
<normalizedToken id="AB907D83A90D24BC24F236ACF0C1A9C2" originalValue="µm">µm</normalizedToken>
, about 3
<normalizedToken id="7B721EBBD72AD36D71E82C200D8787B6" originalValue="µm">µm</normalizedToken>
wide. Clitellum extending over
<normalizedToken id="7DBDB6B59D88DF46CD2916562641633E" originalValue="XII–XIII">XII-XIII</normalizedToken>
.
</paragraph>
<paragraph id="38E0D43A907A847C726DD6C9F8F3E0C8" pageId="5" pageNumber="20">
Coelomocytes 15
<normalizedToken id="B99704E569779088E097307F58CED787" originalValue="µm">µm</normalizedToken>
long. Dorsal vessel originating in XIII. Nephridia observed in
<normalizedToken id="3A1F80D572B48957E7043100A71665ED" originalValue="VIII–X">VIII-X</normalizedToken>
and
<normalizedToken id="1EE005452C2FC8CA98D49285A40FC41C" originalValue="XIII–XX">XIII-XX</normalizedToken>
, about 110
<normalizedToken id="06DAC374B396D463BF9F85B5B40D391A" originalValue="µm">µm</normalizedToken>
long.
</paragraph>
<paragraph id="BF170D82100A88E25DFE3646E4D487AF" pageId="5" pageNumber="20">
Testes originating in XI, extending forwards into X. Sperm funnels folded, length and width unclear. Vasa deferentia 15
<normalizedToken id="7F05CF8478A068E239DAFC8D394F8A31" originalValue="µm">µm</normalizedToken>
wide. Penial bulbs 115
<normalizedToken id="5D5AD7099527538A7782805C45D838BE" originalValue="µm">µm</normalizedToken>
in diameter. No mature eggs observed.
</paragraph>
<paragraph id="78CB86A0EA1EE567DFA5FD9E5D0DD956" pageId="5" pageNumber="20">
Spermathecae (Fig. 2C) 270
<normalizedToken id="3160D2B48EF754BCB8CBC36BAB8211E4" originalValue="µm">µm</normalizedToken>
long, 120
<normalizedToken id="3342C42F7C6D2CFD97D51F3A835F5123" originalValue="µm">µm</normalizedToken>
wide at widest part of ampulla. Glandular body at ectal pore 115
<normalizedToken id="5A5ED10BC4D43562A66170728557D6D2" originalValue="µm">µm</normalizedToken>
in diameter at its widest part. No midventral subneural glands observed.
</paragraph>
</subSubSection>
<subSubSection id="60B94E0AB28B751A73C67BC099018E2E" pageId="5" pageNumber="20" type="distribution">
<paragraph id="CE691D1D7C47A4F59A098FD6A1C48A99" pageId="5" pageNumber="20">Geographical distribution including BOLD data.</paragraph>
<paragraph id="70A256ACE3219165F9E743C7C6192581" pageId="5" pageNumber="20">
Genetically identified from the Netherlands, Norway, Spain and Sweden; also recognized from Canada (BIN-numbers BOLD:AAF9630 &amp; BOLD:ACV7068). This species has historically been widely reported from Europe and North America, and even from the southern hemisphere (
<bibRefCitation id="FE5409B8303CC62A0A414B603DAF79CB" author="Stephenson, J" journalOrPublisher="Discovery Reports" pageId="49" pageNumber="64" pagination="233 - 264" title="Oligochaeta. Part. I. Microdrili." volume="4" year="1932 b">Stephenson 1932b</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="47114946D24B01A104F6D918549C43B5" lastPageId="7" lastPageNumber="22" pageId="5" pageNumber="20" type="remarks">
<paragraph id="8C066A5AA2989C722F7E7984B8280B12" pageId="5" pageNumber="20">Remarks.</paragraph>
<paragraph id="4A7BF918AFC576A3C18BE666BDD09B71" pageId="5" pageNumber="20">
<taxonomicName id="E0D1760A875C8B07F2AB0263BD980B1E" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
, possibly the first ever described enchytraeid, has an interesting history and was for a long time poorly defined as a species. It was given the name
<taxonomicName id="0700E7CCED30B38450D2ECAB77436EE8" class="Clitellata" family="Lumbricidae" genus="Lumbricus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricus lineatus" order="Crassiclitellata" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricus lineatus</taxonomicName>
by
<bibRefCitation id="EC4A2D13DC8624B4A9ED33067EAD493D" pageId="5" pageNumber="20">
<normalizedToken id="84634BE6F588245B288E0170C9103382" originalValue="Müller">Mueller</normalizedToken>
(1774)
</bibRefCitation>
who described it as abundant on the shores of the Baltic Sea.
<normalizedToken id="D5ED7CA564DBDFE3359B7477C913893B" originalValue="Müller">Mueller</normalizedToken>
classified the worm under
<normalizedToken id="BDA938DA9BC47E19C7A6DE86C13783A9" originalValue="“Setosa”">&quot;Setosa&quot;</normalizedToken>
, and gave a brief description of its circulatory system and short protruding chaetae. In the same work, he refers to a worm that he previously described (
<bibRefCitation id="4A933C0A30A366D351EFFCE6DC8C0B3A" pageId="5" pageNumber="20">
<normalizedToken id="42F69D4CBF48D532837683D1C159E824" originalValue="Müller">Mueller</normalizedToken>
1771
</bibRefCitation>
), which he found on sandy shores of the Baltic and particularly among the rotting seaweed by the ramparts of Copenhagen, where he did most of his work. However, he classified that worm in
<taxonomicName id="FA62E9E1CD5032F3D85443D3D0C67751" class="Gordioida" family="Gordiidae" genus="Gordius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gordius" order="Gordioidea" pageId="5" pageNumber="20" phylum="Nematomorpha" rank="genus">Gordius</taxonomicName>
, referred to it as a Faden Wurm (nematode) said to lack any segments or ring, and he failed to mention anything about chaetae. It is therefore difficult to say if
<normalizedToken id="891179A072AF3BE04147E6E20D497E9D" originalValue="Müller">Mueller</normalizedToken>
simply missed those characters or if the worm from 1771 was something different. Nevertheless, as
<normalizedToken id="AFC63F31EB7D5B079F991B59A3055477" originalValue="Müller">Mueller</normalizedToken>
worked with live material, it is most plausible that the type locality for the first described
<taxonomicName id="2EBCC3D8A7EDB6E588F5E77FA5301AF0" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="5" pageNumber="20" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
is around Copenhagen. Unfortunately, we do not have any specimens from Copenhagen, but we have found that our molecularly defined
<taxonomicName id="189A3571F56CCADF9FA6B169EAEB312A" lsidName="L. lineatus" pageId="5" pageNumber="20" rank="species" species="lineatus">L. lineatus</taxonomicName>
is common throughout the Baltic Sea. For the sake of finally defining the true
<taxonomicName id="DA3769C558C8F3E2CF93CD20E0DF371C" lsidName="L. lineatus" pageId="5" pageNumber="20" rank="species" species="lineatus">L. lineatus</taxonomicName>
and connecting it to a molecular profile, we decided to designate one of our specimens from
<normalizedToken id="7A9983FB302831EF29048D5300389A39" originalValue="Öland">Oeland</normalizedToken>
in the Baltic Sea as a neotype.
</paragraph>
<paragraph id="3CF5418B057516CC7FFF6254DDF2289A" lastPageId="7" lastPageNumber="22" pageId="5" pageNumber="20">
The specimens of
<taxonomicName id="E0D6C7AC65E8A7719DD261BBCA783779" lsidName="L. lineatus" pageId="5" pageNumber="20" rank="species" species="lineatus">L. lineatus</taxonomicName>
in this study were smaller than the ones in the re-description by
<bibRefCitation id="13A98631BCFB52346727ADC046AA94B1" pageId="5" pageNumber="20">Nielsen and Christensen (1959)</bibRefCitation>
and the sperm funnels were shorter in relation to their length. However, the shape of the spermathecae, number of chaetae and segments conformed well with these
<normalizedToken id="67866AB7BA9AADC3D60EAE727FF89D29" originalValue="authors’">authors'</normalizedToken>
description. Furthermore, both the vesiculate and avesiculate forms of the species were observed, as has previously been noted to occur in
<taxonomicName id="964FAA3918157BB27D1E0EE1B6323B52" lsidName="L. lineatus" pageId="5" pageNumber="20" rank="species" species="lineatus">L. lineatus</taxonomicName>
(
<bibRefCitation id="A056CD17D60C6504C45C200D62B45FC1" pageId="5" pageNumber="20">Nielsen and Christensen 1959</bibRefCitation>
,
<bibRefCitation id="3864D651B1A7624D3EC48BCCAA96B0C6" author="Christensen, B" journalOrPublisher="Chromosoma" pageId="44" pageNumber="59" url="https://doi.org/10.1111/j.1601-5223.1961.tb01782.x" year="1961">Christensen 1961</bibRefCitation>
), further supporting the
<pageBreakToken id="9596FA6A3BED8014343DD4FA41D79595" pageId="6" pageNumber="21" start="start">designation</pageBreakToken>
of a neotype from our material.
<bibRefCitation id="84BE37C9A37C5EC823C9A01E2E8A046A" author="Christensen, B" journalOrPublisher="Nature" pageId="45" pageNumber="60" pagination="1085 - 1086" title="Pseudofertlization in the genus Lumbricillus (Enchytraeidae)." volume="181" year="1958">
Christensen and
<normalizedToken id="1E31811AF7707582F407398EE9CA9DD1" originalValue="O’Connor">O'Connor</normalizedToken>
(1958)
</bibRefCitation>
were the first to describe diploid vesiculate and triploid avesiculate forms of
<taxonomicName id="DEB78238E95B46969ADB43708C4B41DD" lsidName="L. lineatus" pageId="6" pageNumber="21" rank="species" species="lineatus">L. lineatus</taxonomicName>
and their intriguing life histories (see also
<bibRefCitation id="F00BEC9149DABE5D5DD96EDEDCCA218D" author="Christensen, B" journalOrPublisher="Chromosoma" pageId="44" pageNumber="59" pagination="365 - 379" title="A comparative cytological investigation of the reproductive cycle of an amphimictic diploid and a parthenogenetic triploid form of Lumbricilluslineatus (O. F. M.) (Oligochaeta, Enchytraeidae)." url="https://doi.org/10.1007/BF00328661" volume="11" year="1960">Christensen 1960</bibRefCitation>
). While the diploid form has normal gametogenesis and is amphimictic, the triploid form is dependent on copulation with the diploid form, which acts as sperm donor, to produce offspring. Furthermore, the sperm from the diploid form does merely activate the egg of the triploid individual, without fertilizing it. Instead, oogenesis in the triploid cytotype follows a peculiar pattern of chromosome divisions and mergings that results ultimately in the restoration of triploid nuclei. The relationship between the two forms has been described as an obligatory co-existence (
<bibRefCitation id="A84CB1881D64E7E5A7EC6BA6043E26C0" author="Christensen, B" journalOrPublisher="Chromosoma" pageId="44" pageNumber="59" pagination="365 - 379" title="A comparative cytological investigation of the reproductive cycle of an amphimictic diploid and a parthenogenetic triploid form of Lumbricilluslineatus (O. F. M.) (Oligochaeta, Enchytraeidae)." url="https://doi.org/10.1007/BF00328661" volume="11" year="1960">Christensen 1960</bibRefCitation>
). To further complicate things, tetra- and pentaploids have also been observed within
<taxonomicName id="3512C9F8AD0DFDB7016566CD362A558D" lsidName="L. lineatus" pageId="6" pageNumber="21" rank="species" species="lineatus">L. lineatus</taxonomicName>
. The tetra- and pentaploids have testis sacs that are smaller than those of diploids but larger than those of triploids. Furthermore, the sperm funnels are smaller in tetra- pentaploids than in di- and triploids. Apparently, the testes sacs of tetra- and pentaploids sometimes produce sperm which can activate the eggs of all polyploid
<pageBreakToken id="B298CF3EC8A93FCEE926462F9C298210" pageId="7" pageNumber="22" start="start">forms</pageBreakToken>
(
<bibRefCitation id="B1944F563308BB113AC7A906F7D03A7B" author="Christensen, B" journalOrPublisher="Hereditas" pageId="45" pageNumber="60" pagination="65 - 73" title="Long-term isozyme variation in parthenogenetic polyploid forms of Lumbricilluslineatus (Enchytraeidae, Oligochaeta) in recently established environments." url="https://doi.org/10.1111/j.1601-5223.1978.tb01604.x" volume="88" year="1978">Christensen et al. 1978</bibRefCitation>
). Unfortunately, we did not determine the ploidy level of our sampled specimens, but we did not observe any genetic distinction between the vesiculate and avesiculate forms, neither in COI nor in ITS sequences (Klinth at al. 2017).
</paragraph>
<paragraph id="E9E87389488B93F1E99FA987308B3F1A" pageId="7" pageNumber="22">
<taxonomicName id="30BD49535F144DDED1535654090E5239" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="7" pageNumber="22" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
is morphologically most similar to
<taxonomicName id="F9DA84D1FE7E5CE38CD41D7FA27118CB" lsidName="L. verrucosus" pageId="7" pageNumber="22" rank="species" species="verrucosus">L. verrucosus</taxonomicName>
and
<taxonomicName id="4B13457A8AE48A0E9926258BF2CA1545" lsidName="L. latithecatus" pageId="7" pageNumber="22" rank="species" species="latithecatus">L. latithecatus</taxonomicName>
sp. n. (compare spermathecae in Fig. 3 and see Remarks for these species), but also superficially similar to the other members of the group we have chosen to call the linaetus group. Genetically it is closely related to
<taxonomicName id="2C75D1BD16C48D49954CE2ABC74FBF04" lsidName="L. rutilus" pageId="7" pageNumber="22" rank="species" species="rutilus">L. rutilus</taxonomicName>
Welch, 1914 and
<taxonomicName id="3B011CBE67EC6E88831585209A6427A2" lsidName="L. latithecatus" pageId="7" pageNumber="22" rank="species" species="latithecatus">L. latithecatus</taxonomicName>
sp. n. (
<bibRefCitation id="E796C9DFDB71A8D0B5D4D1029DC13F96" 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>
; where
<taxonomicName id="D9C07D3F7E1C9C949B9D31CEFACC803B" lsidName="L. latithecatus" pageId="7" pageNumber="22" rank="species" species="latithecatus">L. latithecatus</taxonomicName>
is called
<taxonomicName id="657EDF887E0E5F2E9FC30AEA060A2CE0" genus="L." lsidName="L." pageId="7" pageNumber="22" rank="genus">L.</taxonomicName>
sp. E) (Fig. 1).
</paragraph>
<caption id="057369245C19BEA47CE8A92502DAFD81" pageId="7" pageNumber="22">
<paragraph id="F50F194E2D4BD2BA57DB3437B6EA2630" pageId="7" pageNumber="22">
Figure 3. A
<taxonomicName id="DB8497228BEB43B4E68A82F45200A141" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus lineatus" order="Enchytraeida" pageId="7" pageNumber="22" phylum="Annelida" rank="species" species="lineatus">Lumbricillus lineatus</taxonomicName>
, spermatheca B
<taxonomicName id="B52A9ACED808B62921B7D644CC2ABC70" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus latithecatus" order="Enchytraeida" pageId="7" pageNumber="22" phylum="Annelida" rank="species" species="latithecatus">Lumbricillus latithecatus</taxonomicName>
sp. n. (holotype), spermathecae C
<taxonomicName id="683BA27255A6E0165EBC82787E6C13B5" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="7" pageNumber="22" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
, spermatheca D
<taxonomicName id="4EBE17CE34E1124DEEF56D46785B6AF0" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Lumbricillus verrucosus" order="Enchytraeida" pageId="7" pageNumber="22" phylum="Annelida" rank="species" species="verrucosus">Lumbricillus verrucosus</taxonomicName>
, spermatheca seen from below. Scale bars: 100
<normalizedToken id="60ACE52F5B07CB50B9EAFE050AEC8459" originalValue="µm">µm</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>
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